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Article ; Online: Modeling Mammary Organogenesis from Biological First Principles: A Systems Biology Approach.

Schaeberle, Cheryl M / Bouffard, Victoria A / Sonnenschein, Carlos / Soto, Ana M

Methods in molecular biology (Clifton, N.J.)

2023 Volume 2745, Page(s) 177–188

Abstract: Stromal-epithelial interactions mediate mammary gland development and the formation and progression of breast cancer. To study these interactions in vitro, 3D models are essential. We have successfully developed novel 3D in vitro models that allow the ... ...

Abstract	Stromal-epithelial interactions mediate mammary gland development and the formation and progression of breast cancer. To study these interactions in vitro, 3D models are essential. We have successfully developed novel 3D in vitro models that allow the formation of mammary gland structures closely resembling those found in vivo and that respond to the hormonal cues that regulate mammary gland morphogenesis and function. Due to their simplicity when compared to in vivo studies, and to their accessibility to visualization in real time, these models are well suited to conceptual and mathematical modeling.
MeSH term(s)	Humans ; Animals ; Female ; Breast ; Organogenesis/physiology ; Breast Neoplasms ; Mammary Glands, Animal ; Morphogenesis/physiology ; Epithelial Cells
Language	English
Publishing date	2023-12-07
Publishing country	United States
Document type	Journal Article
ISSN	1940-6029
ISSN (online)	1940-6029
DOI	10.1007/978-1-0716-3577-3_11
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Article ; Online: From Wingspread to CLARITY: a personal trajectory.

Soto, Ana M / Schaeberle, Cheryl M / Sonnenschein, Carlos

Nature reviews. Endocrinology

2021 Volume 17, Issue 4, Page(s) 247–256

Abstract: In the three decades since endocrine disruption was conceptualized at the Wingspread Conference, we have witnessed the growth of this multidisciplinary field and the accumulation of evidence showing the deleterious health effects of endocrine-disrupting ... ...

Abstract	In the three decades since endocrine disruption was conceptualized at the Wingspread Conference, we have witnessed the growth of this multidisciplinary field and the accumulation of evidence showing the deleterious health effects of endocrine-disrupting chemicals. It is only within the past decade that, albeit slowly, some changes regarding regulatory measures have taken place. In this Perspective, we address some historical points regarding the advent of the endocrine disruption field and the conceptual changes that endocrine disruption brought about. We also provide our personal recollection of the events triggered by our serendipitous discovery of oestrogenic activity in plastic, a founder event in the field of endocrine disruption. This recollection ends with the CLARITY study as an example of a discordance between 'science for its own sake' and 'regulatory science' and leads us to offer a perspective that could be summarized by the motto attributed to Ludwig Boltzmann: "Nothing is more practical than a good theory".
MeSH term(s)	Animals ; Endocrine Disruptors/toxicity ; Endocrine System/drug effects ; Environmental Exposure/adverse effects ; Fetal Development/drug effects ; Government Regulation ; Humans
Chemical Substances	Endocrine Disruptors
Language	English
Publishing date	2021-01-29
Publishing country	England
Document type	Journal Article ; Research Support, N.I.H., Extramural ; Review
ZDB-ID	2489381-X
ISSN	1759-5037 ; 1759-5029
ISSN (online)	1759-5037
ISSN	1759-5029
DOI	10.1038/s41574-020-00460-3
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Article: The Case for BPA as an Obesogen: Contributors to the Controversy.

Rubin, Beverly S / Schaeberle, Cheryl M / Soto, Ana M

Frontiers in endocrinology

2019 Volume 10, Page(s) 30

Abstract: Since the inception of the term endocrine disruptor, the idea that the environment is an important determinant of phenotype has motivated researchers to explore the effect of low dose exposure to BPA during organogenesis. The syndrome observed was ... ...

Abstract	Since the inception of the term endocrine disruptor, the idea that the environment is an important determinant of phenotype has motivated researchers to explore the effect of low dose exposure to BPA during organogenesis. The syndrome observed was complex, affecting various endpoints such as reproduction and reproductive tissues, behavior, mammary gland development and carcinogenesis, glucose homeostasis, and obesity. This constellation of impacted endpoints suggests the possibility of complex interactions among the multiple effects of early BPA exposure. One key finding of our rodent studies was alterations of energy and amino-acid metabolism that were detected soon after birth and continued to be present at all time points examined through 6 months of age. The classical manifestations of obesity and associated elements of metabolic disease took a longer time to become apparent. Here we examine the validity of the often-mentioned lack of reproducibility of obesogenic effects of BPA, starting from the known environmental causes of variation, which are diverse and range from the theoretical like the individuation process and the non-monotonicity of the dose-response curve, to the very pragmatic like housing, feed, and time and route of exposure. We then explore environmental conditions that may hinder reproducibility and discuss the effect of confounding factors such as BPA-induced hyperactivity. In spite of all the potential sources of variation, we find that some obesogenic or metabolic effects of BPA are reproducibly observed when study conditions are analogous. We recommend that study authors describe details of their study conditions including the environment, husbandry, and feed. Finally, we show that when experimental conditions are strictly maintained, reproducibility, and stability of the obese phenotype is consistently observed.
Language	English
Publishing date	2019-02-06
Publishing country	Switzerland
Document type	Journal Article ; Review
ZDB-ID	2592084-4
ISSN	1664-2392
ISSN	1664-2392
DOI	10.3389/fendo.2019.00030
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Article ; Online: A Combined Morphometric and Statistical Approach to Assess Nonmonotonicity in the Developing Mammary Gland of Rats in the CLARITY-BPA Study.

Montévil, Maël / Acevedo, Nicole / Schaeberle, Cheryl M / Bharadwaj, Manushree / Fenton, Suzanne E / Soto, Ana M

Environmental health perspectives

2020 Volume 128, Issue 5, Page(s) 57001

Abstract: Background: The Consortium Linking Academic and Regulatory Insights on Bisphenol-A (CLARITY-BPA) is a rare collaboration of guideline-compliant (core) studies and academic hypothesis-based studies to assess the effects of bisphenol A (BPA).: ... ...

Abstract	Background: The Consortium Linking Academic and Regulatory Insights on Bisphenol-A (CLARITY-BPA) is a rare collaboration of guideline-compliant (core) studies and academic hypothesis-based studies to assess the effects of bisphenol A (BPA). Objectives: We aimed to Methods: Sprague-Dawley rats were exposed to BPA, vehicle, or positive control [ethinyl estradiol (EE2)] by oral gavage beginning on gestational day (GD)6 and continuing with direct dosing of the pups after birth. There were two studies: subchronic and chronic. The latter used two exposure regimes, one stopping at postnatal day (PND)21 (stop-dose) the other continuing until tissue harvest (continuous). Glands were harvested at multiple time points; whole mounts and histological specimens were analyzed blinded to treatment. Results: The subchronic study's semiquantitative analysis revealed no significant differences between control and BPA dose groups at PND21, whereas at PND90 there were significant differences between control and the lowest BPA dose and between control and the lowest EE2 dose in animals in estrus. Quantitative, automatized analysis of the chronic PND21 specimens displayed nonmonotonic BPA effects, with a breaking point between the 25 and Conclusions: Both the semiquantitative and the quantitative methods revealed nonmonotonic effects of BPA. The quantitative unsupervised analysis used
MeSH term(s)	Animals ; Benzhydryl Compounds/toxicity ; Ethinyl Estradiol/toxicity ; Female ; Hazardous Substances/toxicity ; Mammary Glands, Animal/drug effects ; Mammary Glands, Animal/growth & development ; Phenols/toxicity ; Pregnancy ; Rats ; Rats, Sprague-Dawley
Chemical Substances	Benzhydryl Compounds ; Hazardous Substances ; Phenols ; Ethinyl Estradiol (423D2T571U) ; bisphenol A (MLT3645I99)
Language	English
Publishing date	2020-05-20
Publishing country	United States
Document type	Journal Article ; Research Support, N.I.H., Extramural ; Research Support, N.I.H., Intramural
ZDB-ID	195189-0
ISSN	1552-9924 ; 0091-6765 ; 1078-0475
ISSN (online)	1552-9924
ISSN	0091-6765 ; 1078-0475
DOI	10.1289/EHP6301
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Article ; Online: Perinatal BPA exposure and reproductive axis function in CD-1 mice.

Acevedo, Nicole / Rubin, Beverly S / Schaeberle, Cheryl M / Soto, Ana M

Reproductive toxicology (Elmsford, N.Y.)

2018 Volume 79, Page(s) 39–46

Abstract: Perinatal Bisphenol-A (BPA) exposure reduces fertility and fecundity in mice. This study examined effects of early BPA exposure on activation of gonadotropin releasing hormone (GnRH) neurons in conjunction with a steroid-induced luteinizing hormone (LH) ... ...

Abstract	Perinatal Bisphenol-A (BPA) exposure reduces fertility and fecundity in mice. This study examined effects of early BPA exposure on activation of gonadotropin releasing hormone (GnRH) neurons in conjunction with a steroid-induced luteinizing hormone (LH) surge, characterized patterns of estrous cyclicity and fertility over time, and assessed the ovarian follicular reserve to further explore factors responsible for the reduced fertility we previously described in this model. The percent activated GnRH neurons was reduced in BPA-exposed females at 3-6 months, and periods of persistent proestrus were increased. These data suggest that perinatal exposure to BPA reduces GnRH neuronal activation required for the generation of the LH surge and estrous cyclicity. Assessments of anti-Müllerian hormone (AMH) levels failed to suggest a decline in the follicular reserve at the BPA exposure levels examined.
MeSH term(s)	Animals ; Anti-Mullerian Hormone/blood ; Benzhydryl Compounds/toxicity ; Endocrine Disruptors/toxicity ; Estrogens/toxicity ; Estrous Cycle/drug effects ; Female ; Fertility/drug effects ; Gonadotropin-Releasing Hormone/metabolism ; Luteinizing Hormone/blood ; Male ; Maternal-Fetal Exchange ; Mice ; Neurons/drug effects ; Neurons/metabolism ; Phenols/toxicity ; Pregnancy
Chemical Substances	Benzhydryl Compounds ; Endocrine Disruptors ; Estrogens ; Phenols ; Gonadotropin-Releasing Hormone (33515-09-2) ; Anti-Mullerian Hormone (80497-65-0) ; Luteinizing Hormone (9002-67-9) ; bisphenol A (MLT3645I99)
Language	English
Publishing date	2018-05-09
Publishing country	United States
Document type	Journal Article ; Research Support, N.I.H., Extramural
ZDB-ID	639342-1
ISSN	1873-1708 ; 0890-6238
ISSN (online)	1873-1708
ISSN	0890-6238
DOI	10.1016/j.reprotox.2018.05.002
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Article ; Online: New insights into fetal mammary gland morphogenesis: differential effects of natural and environmental estrogens.

Speroni, Lucia / Voutilainen, Maria / Mikkola, Marja L / Klager, Skylar A / Schaeberle, Cheryl M / Sonnenschein, Carlos / Soto, Ana M

Scientific reports

2017 Volume 7, Page(s) 40806

Abstract: An increased breast cancer risk during adulthood has been linked to estrogen exposure during fetal life. However, the impossibility of removing estrogens from the feto-maternal unit has hindered the testing of estrogen's direct effect on mammary gland ... ...

Abstract	An increased breast cancer risk during adulthood has been linked to estrogen exposure during fetal life. However, the impossibility of removing estrogens from the feto-maternal unit has hindered the testing of estrogen's direct effect on mammary gland organogenesis. To overcome this limitation, we developed an ex vivo culture method of the mammary gland where the direct action of estrogens can be tested during embryonic days (E)14 to 19. Mouse mammary buds dissected at E14 and cultured for 5 days showed that estrogens directly altered fetal mammary gland development. Exposure to 0.1 pM, 10 pM, and 1 nM 17 β-estradiol (E2) resulted in monotonic inhibition of mammary buds ductal growth. In contrast, Bisphenol-A (BPA) elicited a non-monotonic response. At environmentally relevant doses (1 nM), BPA significantly increased ductal growth, as previously observed in vivo, while 1 μM BPA significantly inhibited ductal growth. Ductal branching followed the same pattern. This effect of BPA was blocked by Fulvestrant, a full estrogen antagonist, while the effect of estradiol was not. This method may be used to study the hormonal regulation of mammary gland development, and to test newly synthesized chemicals that are released into the environment without proper assessment of their hormonal action on critical targets like the mammary gland.
MeSH term(s)	Animals ; Benzhydryl Compounds/pharmacology ; Cell Proliferation/drug effects ; Estradiol/analogs & derivatives ; Estradiol/pharmacology ; Female ; Fetus/cytology ; Fulvestrant ; Mammary Glands, Animal/cytology ; Mammary Glands, Animal/drug effects ; Mammary Glands, Animal/growth & development ; Mammary Glands, Animal/pathology ; Mice ; Morphogenesis/drug effects ; Phenols/pharmacology ; Pregnancy ; Receptors, Estrogen/metabolism ; Receptors, G-Protein-Coupled/metabolism
Chemical Substances	Benzhydryl Compounds ; GPER1 protein, mouse ; Phenols ; Receptors, Estrogen ; Receptors, G-Protein-Coupled ; Fulvestrant (22X328QOC4) ; Estradiol (4TI98Z838E) ; bisphenol A (MLT3645I99)
Language	English
Publishing date	2017-01-19
Publishing country	England
Document type	Journal Article ; Research Support, N.I.H., Extramural ; Research Support, Non-U.S. Gov't
ZDB-ID	2615211-3
ISSN	2045-2322 ; 2045-2322
ISSN (online)	2045-2322
ISSN	2045-2322
DOI	10.1038/srep40806
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Article ; Online: The male mammary gland: a target for the xenoestrogen bisphenol A.

Vandenberg, Laura N / Schaeberle, Cheryl M / Rubin, Beverly S / Sonnenschein, Carlos / Soto, Ana M

Reproductive toxicology (Elmsford, N.Y.)

2013 Volume 37, Page(s) 15–23

Abstract: Males of some strains of mice retain their mammary epithelium even in the absence of nipples. Here, we have characterized the mammary gland in male CD-1 mice both in whole mounts and histological sections. We also examined the effects of bisphenol A (BPA) ...

Abstract	Males of some strains of mice retain their mammary epithelium even in the absence of nipples. Here, we have characterized the mammary gland in male CD-1 mice both in whole mounts and histological sections. We also examined the effects of bisphenol A (BPA), an estrogen mimic that alters development of the female mouse mammary gland. BPA was administered at a range of environmentally relevant doses (0.25-250μg/kg/day) to pregnant and lactating mice and then the mammary glands of male offspring were examined at several periods in adulthood. We observed age- and dose-specific effects on mammary gland morphology, indicating that perinatal BPA exposures alter the male mammary gland in adulthood. These results may provide insight into gynecomastia, the most common male breast disease in humans, where proliferation of the mammary epithelium leads to breast enlargement.
MeSH term(s)	Age Factors ; Animals ; Benzhydryl Compounds/toxicity ; Dose-Response Relationship, Drug ; Epithelium/anatomy & histology ; Epithelium/drug effects ; Estrogens/toxicity ; Female ; Male ; Mammary Glands, Animal/anatomy & histology ; Mammary Glands, Animal/drug effects ; Mice ; Phenols/toxicity ; Pregnancy
Chemical Substances	Benzhydryl Compounds ; Estrogens ; Phenols ; bisphenol A (MLT3645I99)
Language	English
Publishing date	2013-01-21
Publishing country	United States
Document type	Journal Article ; Research Support, N.I.H., Extramural
ZDB-ID	639342-1
ISSN	1873-1708 ; 0890-6238
ISSN (online)	1873-1708
ISSN	0890-6238
DOI	10.1016/j.reprotox.2013.01.002
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Article ; Online: Perinatally administered bisphenol a as a potential mammary gland carcinogen in rats.

Acevedo, Nicole / Davis, Barbara / Schaeberle, Cheryl M / Sonnenschein, Carlos / Soto, Ana M

Environmental health perspectives

2013 Volume 121, Issue 9, Page(s) 1040–1046

Abstract: Background: Environmental exposure to bisphenol A (BPA) affects mammary gland development in rodents and primates. Prenatal exposure to environmentally relevant doses of BPA increased the number of intraductal hyperplasias and ductal carcinomas in situ ... ...

Abstract	Background: Environmental exposure to bisphenol A (BPA) affects mammary gland development in rodents and primates. Prenatal exposure to environmentally relevant doses of BPA increased the number of intraductal hyperplasias and ductal carcinomas in situ by 50 days of age in Wistar-Furth rats. Objective: We aimed to determine whether BPA exposure of dams during gestation only or throughout lactation affects the incidence of mammary gland neoplasia in female offspring. Methods: We treated pregnant Sprague-Dawley rats with BPA at 0, 0.25, 2.5, 25, or 250 μg BPA/kg BW/day from gestational day (GD) 9 to birth and from GD9 to postnatal day (PND) 21. Mammary glands from BPA-exposed offspring were examined at four time points for preneoplastic and neoplastic lesions. To assess circulating BPA levels, we exposed pregnant rats to vehicle or 250 μg BPA/kg BW/day during gestation only or during gestation/lactation and analyzed sera from dams, fetuses, and nursing pups for total and unconjugated BPA. Results: Total and unconjugated BPA were detected in sera from 100% of dams and fetuses and 33% of pups exposed to 250 μg BPA/kg BW/day. Unconjugated BPA levels in exposed dams and fetuses (gestational) and in exposed dams and pups (gestational/lactational) were within levels found in humans. Preneoplastic lesions developed in BPA-exposed female offspring across all doses as early as PND50. Unexpectedly, mammary gland adenocarcinomas developed in BPA-exposed offspring by PND90. Conclusions: Our findings suggest that developmental exposure to environmentally relevant levels of BPA during gestation and lactation induces mammary gland neoplasms in the absence of any additional carcinogenic treatment. Thus, BPA may act as a complete mammary gland carcinogen.
MeSH term(s)	Animals ; Benzhydryl Compounds/administration & dosage ; Benzhydryl Compounds/blood ; Benzhydryl Compounds/toxicity ; Carcinogens, Environmental/administration & dosage ; Carcinogens, Environmental/toxicity ; Carcinoma, Ductal/chemically induced ; Environmental Exposure/adverse effects ; Female ; Hyperplasia/chemically induced ; Mammary Neoplasms, Animal/chemically induced ; Phenols/administration & dosage ; Phenols/blood ; Phenols/toxicity ; Pregnancy ; Rats ; Rats, Inbred WF ; Rats, Sprague-Dawley
Chemical Substances	Benzhydryl Compounds ; Carcinogens, Environmental ; Phenols ; bisphenol A (MLT3645I99)
Language	English
Publishing date	2013-07-23
Publishing country	United States
Document type	Journal Article ; Research Support, N.I.H., Extramural
ZDB-ID	195189-0
ISSN	1552-9924 ; 0091-6765 ; 1078-0475
ISSN (online)	1552-9924
ISSN	0091-6765 ; 1078-0475
DOI	10.1289/ehp.1306734
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Zs.MO 379: Show issues

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Article: Strengths and weaknesses of in vitro assays for estrogenic and androgenic activity.

Soto, Ana M / Maffini, Maricel V / Schaeberle, Cheryl M / Sonnenschein, Carlos

Best practice & research. Clinical endocrinology & metabolism

2006 Volume 20, Issue 1, Page(s) 15–33

Abstract: The endocrine and reproductive effects of xenobiotics are believed to be due to (1) their mimicking the effects of endogenous hormones; (2) their antagonizing the effects of endogenous hormones; (3) their altering the pattern of synthesis and metabolism ... ...

Abstract	The endocrine and reproductive effects of xenobiotics are believed to be due to (1) their mimicking the effects of endogenous hormones; (2) their antagonizing the effects of endogenous hormones; (3) their altering the pattern of synthesis and metabolism of natural hormones; and (4) their modifying hormone receptor levels. It has been suggested that endocrine disruptors may play a role in the decrease in human semen quantity and quality, an increase in the anomalies of male genital tract, and an increase in the testicular and breast cancer incidence during the last 50 years. Testing these hypotheses will require: (1) identifying estrogen and androgen agonists and antagonists among the chemicals present in the environment; (2) assessing the interactions among the endocrine disruptors to which humans are exposed; and (3) finding markers of estrogen (and androgen) exposure. The development of fast and sensitive bioassays is central to the achievement of these three goals.
MeSH term(s)	Androgen Antagonists/pharmacology ; Androgens/agonists ; Androgens/analysis ; Animals ; Binding, Competitive ; Biological Assay/standards ; Breast Neoplasms ; Cell Line ; Cell Line, Tumor ; Cell Proliferation/drug effects ; Dose-Response Relationship, Drug ; Drug Evaluation, Preclinical/methods ; Drug Evaluation, Preclinical/standards ; Endocrine Disruptors/analysis ; Estrogen Antagonists/pharmacology ; Estrogen Receptor alpha/physiology ; Estrogens/analysis ; Female ; Genes, Reporter/physiology ; Humans ; Male ; Receptors, Estrogen/agonists
Chemical Substances	Androgen Antagonists ; Androgens ; Endocrine Disruptors ; Estrogen Antagonists ; Estrogen Receptor alpha ; Estrogens ; Receptors, Estrogen
Language	English
Publishing date	2006-03
Publishing country	Netherlands
Document type	Comparative Study ; Journal Article ; Research Support, N.I.H., Extramural ; Research Support, Non-U.S. Gov't ; Research Support, U.S. Gov't, Non-P.H.S. ; Review
ZDB-ID	2052339-7
ISSN	1532-1908 ; 1521-690X
ISSN (online)	1532-1908
ISSN	1521-690X
DOI	10.1016/j.beem.2005.09.001
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Article ; Online: Dynamic Metabolic Disruption in Rats Perinatally Exposed to Low Doses of Bisphenol-A.

Tremblay-Franco, Marie / Cabaton, Nicolas J / Canlet, Cécile / Gautier, Roselyne / Schaeberle, Cheryl M / Jourdan, Fabien / Sonnenschein, Carlos / Vinson, Florence / Soto, Ana M / Zalko, Daniel

PloS one

2015 Volume 10, Issue 10, Page(s) e0141698

Abstract: Along with the well-established effects on fertility and fecundity, perinatal exposure to endocrine disrupting chemicals, and notably to xeno-estrogens, is strongly suspected of modulating general metabolism. The metabolism of a perinatally exposed ... ...

Abstract	Along with the well-established effects on fertility and fecundity, perinatal exposure to endocrine disrupting chemicals, and notably to xeno-estrogens, is strongly suspected of modulating general metabolism. The metabolism of a perinatally exposed individual may be durably altered leading to a higher susceptibility of developing metabolic disorders such as obesity and diabetes; however, experimental designs involving the long term study of these dynamic changes in the metabolome raise novel challenges. 1H-NMR-based metabolomics was applied to study the effects of bisphenol-A (BPA, 0; 0.25; 2.5, 25 and 250 μg/kg BW/day) in rats exposed perinatally. Serum and liver samples of exposed animals were analyzed on days 21, 50, 90, 140 and 200 in order to explore whether maternal exposure to BPA alters metabolism. Partial Least Squares-Discriminant Analysis (PLS-DA) was independently applied to each time point, demonstrating a significant pair-wise discrimination for liver as well as serum samples at all time-points, and highlighting unequivocal metabolic shifts in rats perinatally exposed to BPA, including those exposed to lower doses. In BPA exposed animals, metabolism of glucose, lactate and fatty acids was modified over time. To further explore dynamic variation, ANOVA-Simultaneous Component Analysis (A-SCA) was used to separate data into blocks corresponding to the different sources of variation (Time, Dose and Time*Dose interaction). A-SCA enabled the demonstration of a dynamic, time/age dependent shift of serum metabolome throughout the rats' lifetimes. Variables responsible for the discrimination between groups clearly indicate that BPA modulates energy metabolism, and suggest alterations of neurotransmitter signaling, the latter finding being compatible with the neurodevelopmental effect of this xenoestrogen. In conclusion, long lasting metabolic effects of BPA could be characterized over 200 days, despite physiological (and thus metabolic) changes connected with sexual maturation and aging.
MeSH term(s)	Animals ; Benzhydryl Compounds/administration & dosage ; Benzhydryl Compounds/pharmacology ; Energy Metabolism/drug effects ; Estrogens, Non-Steroidal/administration & dosage ; Estrogens, Non-Steroidal/pharmacology ; Female ; Liver/drug effects ; Liver/metabolism ; Male ; Metabolome/drug effects ; Phenols/administration & dosage ; Phenols/pharmacology ; Pregnancy ; Proton Magnetic Resonance Spectroscopy/methods ; Rats
Chemical Substances	Benzhydryl Compounds ; Estrogens, Non-Steroidal ; Phenols ; bisphenol A (MLT3645I99)
Language	English
Publishing date	2015-10-30
Publishing country	United States
Document type	Journal Article ; Research Support, N.I.H., Extramural
ISSN	1932-6203
ISSN (online)	1932-6203
DOI	10.1371/journal.pone.0141698
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