Neuroscience of sex differences
Neuroscience of sex differences is the study of understanding the characteristics of the brain that separate the male brain and the female brain. Psychological sex differences are thought by some to reflect the interaction of genes, hormones and social learning on brain development throughout the lifespan.
Some evidence from brain morphology and function studies indicates that male and female brains cannot always be assumed to be identical from either a structural or functional perspective, and some brain structures are sexually dimorphic.
Ideas of differences in the male and female brain circulated during the time of ancient Greek philosophers around 850 B.C. Aristotle claimed that males did not "receive their soul" until 40 days post-gestation and females did not until 80 days. In 1854, Emil Huschke discovered that "the frontal lobe in the male is all of 1% larger than that of the female." As the 19th century progressed, scientists began researching sexual dimorphisms in the brain significantly more. Until around 21 years ago, scientists knew of several structural sexual dimorphisms of the brain, but they did not think that sex had any impact on how the human brain performs daily tasks. Through molecular, animal, and neuroimaging studies, a great deal of information regarding the differences between male and female brains and how much they differ in regards to both structure and function has been uncovered.
Females show enhanced information recall compared to males. This may be due to the fact that females have a more intricate evaluation of risk-scenario contemplation, based on a prefrontal cortical control of the amygdala. For example, the ability to recall information better than males most likely originated from sexual selective pressures on females during competition with other females in mate selection. Recognition of social cues was an advantageous characteristic because it ultimately maximized offspring and was therefore selected for during evolution.
Oxytocin is a hormone that induces contraction of the uterus and lactation in mammals. It is also a characteristic hormone of nursing mothers. Studies have found that oxytocin improves spatial memory. Through activation of the MAP kinase pathway, oxytocin plays a role in the enhancement of long-term synaptic plasticity, which is a change in strength between two neurons over a synapse that lasts for minutes or longer, and long-term memory. This hormone may have helped mothers remember the location of distant food sources so they could better nurture their offspring.
Male vs. female brain anatomyEdit
A popular theory regarding language functions suggests that women use both hemispheres more equally, whereas men are more strongly lateralized to the left hemisphere; however a 2008 meta-analysis of 29 studies comparing language lateralization in males and females found no overall difference.
A 2016 meta-analysis found that the amygdala is not significantly larger in either sex. A 2008 meta-analysis of fMRI studies of amygdala activation found larger effect sizes in men compared with women.
Amygdala volume correlates positively with fearfulness in girls but not in boys.
In fMRI studies, women display stronger activation in the amygdala and prefrontal cortex than men.
A 2016 meta-analysis found that the hippocampus does not differ in volume between men and women.
Other regions and not region-specificEdit
According to one study, there are sex differences in locus coeruleus dendritic structure that allow for an increased reception and processing of limbic information in females compared to males. More research is needed. 
In a 2013 meta-analysis, researchers found on average males had larger grey matter (GM) volume in bilateral amygdalae, hippocampi, anterior parahippocampal gyri, posterior cingulate gyri, precuneus, putamen and temporal poles, areas in the left posterior and anterior cingulate gyri, and areas in the cerebellum bilateral VIIb, VIIIa and Crus I lobes, left VI and right Crus II lobes.  On the other hand, females on average had larger grey matter volume at the right frontal pole, inferior and middle frontal gyri, pars triangularis, planum temporale/parietal operculum, anterior cingulate gyrus, insular cortex, and Heschl's gyrus; bilateral thalami and precuneus; the left parahippocampal gyrus and lateral occipital cortex (superior division). The meta-analysis found larger volumes in females were most pronounced in areas in the right hemisphere related to language in addition to several limbic structures such as the right insular cortex and anterior cingulate gyrus.
Researchers found that female-to-male transgender individuals, sexually attracted to females, who have not undergone surgery or hormonal treatments have white matter microstructure of the superior longitudinal fascicles and forceps minor that is more like cismales than cisfemales. Dimorphism of the putamen may depend not only on genetic sex and experienced gender, but also on sexual orientation, because three studies of transgender populations (all without transition surgery or hormonal treatments) with varying sexual orientation demonstrated varying dimorphism of the putamen. Researchers found using magnetic resonance imaging (MRI) that male-to-female transgender individuals of mixed sexual orientation have a right putamen volume that is significantly different than cismales, and not different than cisfemales. Researchers also found that male-to-female transgender individuals, sexually attracted to females, have a smaller putamen than both cisgender individuals. Researchers also found that male-to-female transgender individuals, sexually attracted to males, have no significant differences in right putamen volume from either cisgender, but female-to-male transgender individuals, sexually attracted to females, like cismales, have larger right putamen than cisfemales.
Male vs. female brain functionEdit
Researchers have found female-typical hypothalamic activation in male-to-female transgender individuals when exposed to one odorous steroid and male-typical activation with exposure to a different odorous steroid. Male-to-female and female-to-male transgender individuals who have not undergone any surgical or hormonal treatment are more likely to be left-handed than cisgender individuals. Male-to-female transgender children, ages 3-12, who have not undergone any surgical or hormonal treatment are more often left-handed than cisgender males, and cisgender males are more likely to be left-handed than cisgender females.
A 2014 meta-analysis found that although men and women commonly used the same brain networks for working memory, specific regions were sex specific. For example, both men and women's active working memory networks composed of bilateral middle frontal gyri, left cingulate gyrus, right precuneus, left inferior and superior parietal lobes, right claustrum, and left middle temporal gyrus but women also tended have consistent activity in the limbic regions such as the anterior cingulate, bilateral amygdala and right hippocampus while men tended to have a distributed networks spread out among the cerebellum, portions of the superior parietal lobe, the left insula and bilateral thalamus.
A 2017 review from the perspective of large-scale brain networks, hypothesized that women's higher susceptibility to stress-prone diseases like PTSD and major depressive disorder, in which the salience network is theorized to be overactive and to interfere with the executive control network, may be due in part (along with societal exposure to stressors and the coping strategies that are available to women) to underlying sex-based brain differences.
Steroid hormones have several effects on brain development as well as maintenance of homeostasis throughout adulthood. One effect they exhibit is on the hypothalamus, where they increase synapse formation. Estrogen receptors have been found in the hypothalamus, pituitary gland, hippocampus, and frontal cortex, indicating the estrogen plays a role in brain development. Gonadal hormone receptors have also been found in the basal fore-brain nuclei.
Estrogen and the female brainEdit
Estradiol influences cognitive function, specifically by enhancing learning and memory in a dose-sensitive manner. Too much estrogen can have negative effects by weakening performance of learned tasks as well as hindering performance of memory tasks; this can result in females exhibiting poorer performance of such tasks when compared to males.
It has been suggested that during development, estrogen can exhibit both feminizing and defeminizing effects on the human brain; high levels of estrogen induce male neural traits to develop while moderate levels induce female traits. In females, defeminizing effects are resisted because of the presence of α-fetoprotein (AFP), a carrier protein proposed to transport estrogen into brain cells, allowing the female brain to properly develop. The role of AFP is significant at crucial stages of development, however. Prenatally, AFP blocks estrogen. Postnatally, AFP decreases to ineffective levels; therefore, it is probable that estrogen exhibits its effects on female brain development postnatally.
Ovariectomies, surgeries inducing menopause, or natural menopause cause fluctuating and decreased estrogen levels in women. This in turn can "attenuate the effects" of endogenous opioid peptides. Opioid peptides are known to play a role in emotion and motivation. β-endorphin (β-EP), an endogenous opioid peptide, content has been found to decrease (in varying amounts/brain region), post ovariectomy, in female rats within the hypothalamus, hippocampus, and pituitary gland. Such a change in β-EP levels could be the cause of mood swings, behavioral disturbances, and hot flashes in post menopausal women.
Testosterone and the male brainEdit
Testosterone heavily influences male development; a study found that perinatal females introduced to elevated testosterone levels exhibited male behavior patterns. In the absence of testosterone, female behavior is retained. Testosterone's influence on the brain is caused by organizational developmental effects. It has been shown to influence proaptotic proteins so that they increase neuronal cell death in certain brain regions. Another way testosterone affects brain development is by aiding in the construction of the "limbic hypothalamic neural networks".
Oxytocin and VasopressinEdit
Oxytocin is positively correlated with maternal behaviours, social recognition, social contact, sexual behaviour and pair bonding. Oxytocin appears at higher levels in women than in men. Vasopressin on the other hand is more present in men and mediates sexual behavior, aggression and other social functions.
It was once thought that sex differences in cognitive task and problem solving did not occur until puberty. However, as of 2000 evidence suggested that cognitive and skill differences are present earlier in development. For example, researchers have found that three- and four-year-old boys were better at targeting and at mentally rotating figures within a clock face than girls of the same age were. Prepubescent girls, however, excelled at recalling lists of words. These sex differences in cognition correspond to patterns of ability rather than overall intelligence. Laboratory settings are used to systematically study the sexual dimorphism in problem solving task performed by adults.
On average, males excel relative to females at certain spatial tasks. Specifically, males have an advantage in tests that require the mental rotation or manipulation of an object. In a computer simulation of a maze task, males completed the task faster and with fewer errors than their female counterparts. Additionally, males have displayed higher accuracy in tests of targeted motor skills, such as guiding projectiles. Males are also faster on reaction time and finger tapping tests.
On average, females excel relative to males on tests that measure recollection. They have an advantage on processing speed involving letters, digits and rapid naming tasks. Females tend to have better object location memory and verbal memory. They also perform better at verbal learning. Females have better performance at matching items and precision tasks, such as placing pegs into designated holes. In maze and path completion tasks, males learn the goal route in fewer trials than females, but females remember more of the landmarks presented. This shows that females use landmarks in everyday situations to orient themselves more than males. Females are better at remembering whether objects had switched places or not.
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