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This article says

"Sex is based on biological factors such as sex chromosomes and gonads [reproductive organs]," she said, "whereas gender has a social component and involves expectations and behaviors based on an individual's perceived sex."

These behaviors and expectations around gender identity can be seen in "epigenetic marks" in the brain, which drive biological functions and features as diverse as memory, development and disease susceptibility. Forger explained that epigenetic marks help determine which genes are expressed and are sometimes passed on from cell to cell as they divide. They also can be passed down from one generation to the next, she said.

"While we are accustomed to thinking about differences between the brains of males and females, we are much less used to thinking about the biological implications of gender identity," she said. "There is now sufficient evidence to suggest that an epigenetic imprint for gender is a logical conclusion. It would be strange if this were not the case, because all environmental influences of any importance can epigenetically change the brain."

This is interesting as I thought gender identity is inborn as seen here

So are you born this way, or a product of the environment?

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What gender someone identifies with or relates to comes in to play outside of utero as kids start to perceive themselves in a certain way. Biological factors, however, influence the likelihood someone will perceive themselves in a certain way. These factors may appear in utero or even later in life. Studies have suggested that developmental factors that occur in utero are likely predictors for the neurological correlates of transsexualism. Transsexualism typically becomes apparent, although in varying degrees of presentation, early on in childhood. It should also be considered that social factors also lead to gendered categorization of what’s considered typical of a particular sex which may also influence the behavior of an individual.

"There are studies suggesting that the development of transsexualism is related to biological mechanisms, such as genetic factors and prenatal exposure to sex hormones, rather than environmental factors like parenting practices. During pregnancy, the foetal brain develops into a male brain under the influence of the androgen testosterone, and into a female brain in the absence of this hormone. The sexual differentiation of the brain however, occurs much later in development than that of the genitals. These two processes could be influenced independently, which is postulated to result in transsexualism if they develop in opposing directions. Thus, transsexualism is believed to result from a discrepancy between sexual brain and genital differentiation caused by genetic or hormonal deviations...

As the research on transsexualism is predominantly carried out on adults it is unclear whether the observed findings have developed early or later in development or before the onset of overt gender dysphoric symptoms. It would be important to also collect structural and functional data from children and adolescents with gender dysphoric symptoms to determine when neural differences between transsexual and non-transsexual individuals emerge." (Smith et al. 2015)

"For children, cross-gender behaviors may start between ages 2 and 4, the same age at which most typically developing children begin showing gendered behaviors and interests. Gender atypical behavior is common among young children and may be part of normal development. Children who meet the criteria for gender dysphoria may or may not continue to experience it into adolescence and adulthood. Some research shows that children who had more intense symptoms and distress, who were more persistent, insistent and consistent in their cross-gender statements and behaviors, and who used more declarative statements (“I am a boy (or girl)” rather than “I want to be a boy (or girl)”) were more likely to become transgender adults." (APA)

"The gonads differentiate after the first 10 weeks of fetal life in humans; thereafter, sex differences in gonadal hormones can have acute or lasting effects on the epigenome. The gendered experiences described in this review start as early as birth (early life stress based on gender) and continue into adolescence and adulthood (cosmetic use, alcohol consumption). Many other gendered experiences not explicitly addressed in this review will also impact the neuroepigenome. The relative contribution of various factors and how they may change throughout development are not known, but the effects of biological sex and gender will interact in myriad ways throughout life. In some cases, gender may amplify epigenetic differences due to sex, whereas in other cases, gendered experiences may counteract differences in the epigenome based on biological sex. Not shown here is the fact that with our current ability to know the sex of an unborn child, gender can start before birth." (Cortes et al. 2019)
(Note: This quote comes from the cited literature in the first article you referenced in your question.)

Sources & Further Reading

Altinay, M., & Anand, A. (2019). Neuroimaging gender dysphoria: a novel psychobiological model. Brain Imaging and Behavior. doi:10.1007/s11682-019-00121-8

PubMed | Google Scholar | Crossref

Cortes, L. R., Cisternas, C. D., & Forger, N. G. (2019). Does Gender Leave an Epigenetic Imprint on the Brain? Frontiers in Neuroscience, 13. doi:10.3389/fnins.2019.00173

PubMed | Google Scholar | Crossref

Manzouri, A., & Savic, I. (2018). Possible Neurobiological Underpinnings of Homosexuality and Gender Dysphoria. Cerebral Cortex, 29(5), 2084-2101. doi:10.1093/cercor/bhy090

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Williams SM, McNamara JO, LaMantia A, Katz LC, Fitzpatrick D, Augustine GJ, Purves D (2001). "What Is Sex?". Neuroscience (2nd ed.). Sunderland (MA): Sinauer Associates. ISBN 978-0-87893-742-4.

PubMed

Schiebinger L, Stefanick ML. Gender matters in biological research and medical practice. J Am Coll Cardiol. (2016);67(2):136-138.

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Smith, E. S., Junger, J., Derntl, B., & Habel, U. (2015). The transsexual brain – A review of findings on the neural basis of transsexualism. Neuroscience & Biobehavioral Reviews, 59, 251-266. doi:10.1016/j.neubiorev.2015.09.008

PubMed | Google Scholar | Crossref

The American Psychiatric Association. What Is Gender Dysphoria?

Webpage

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