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Actin filaments, microtubules MTsand homo filaments, as well as the proteins associated with them, homo a major homo in synapse and dendritic spine formation 2. This implies that the actions of sex hormones in neuronal homo are the result of adrenal, gonadal, and brain local synthesis.
No use, distribution or reproduction Pina sex permitted which does not comply with these terms. This article has been cited by other articles seex PMC. Abstract Piina the brain of female mammals, including humans, a number of physiological and behavioral changes occur as a result of sex hormone exposure. Estradiol and progesterone regulate several brain functions, including Pnia and memory. Sex hormones contribute to Pin the central nervous system by modulating the formation and turnover of the interconnections between neurons as well as controlling the function of glial cells. The dynamics of neuron and glial cells morphology depends on the cytoskeleton and its associated proteins.
Cytoskeletal proteins are necessary to form neuronal dendrites and dendritic spines, as well as to regulate the diverse functions in astrocytes. The expression pattern of proteins, such as actin, microtubule-associated protein 2, Tau, and glial fibrillary acidic protein, changes in a tissue-specific manner in the brain, particularly when variations in sex hormone levels occur during the estrous or menstrual cycles or pregnancy. Here, we review the changes in structure and organization of neurons and glial cells that require the participation of cytoskeletal proteins whose expression and activity are regulated by estradiol and progesterone.
Structural plasticity is highly involved in the functional adaptation of the CNS in response to different environmental and physiological eex, including changes in hormone levels. In particular, female sex hormones can modify the size, morphology, synaptic density, and function of neuronal cells as well as the morphology of glial cells in sex steroid-responsive structures of the CNS 1. These changes are due to modifications in the neuronal and glial cytoskeleton where intracellular signals converge to regulate the direction and speed of outgrowth of different cell structures.
Actin filaments, microtubules MTsand intermediate filaments, as well as the proteins associated with them, play a major role in synapse and dendritic spine formation 2.
Neuronal projections are not only dependent on Pnia activity but also reliant on glial cells. The glia has an essential role in regulating the activity of CNS, where a mutual communication between glial cells and neurons exists. The activity and modifications in glial cell morphology also affect the formation and sxe of synaptic contacts 3Pins. In this review, we will focus on the effects of female sex hormones on the expression and regulation of cytoskeletal proteins, contributing to the remodeling of the adult brain. Sex Hormones and the Brain Female sex hormones are known to have a wide range of effects in the brain regulating not only reproductive processes but also cognitive functions.
Estradiol E2 and progesterone P4 are cholesterol-derived hormones that, given their lipophilic structure, xex easily cross the blood—brain barrier and interact with their specific receptors in different target cells of the brain. These hormones are also synthesized inside the brain. P4 and E2 levels have been detected in different brain areas such se hypothalamus and hippocampus with concentration differences between female and male animals 5 — 7and their synthesis in neurons and glial cells Pinaa been demonstrated 89. Moreover, pregnenolone, a cholesterol metabolite used by neurons for the biosynthesis seex P4 and E2, is also produced by the glia 10 This implies that the actions of sex hormones in neuronal plasticity are the result of adrenal, gonadal, and brain local synthesis.
E2 and P4 effects depend on the signaling pathway they activate, which can be either through intracellular receptors classical mechanism or membrane receptors nonclassical mechanism Female sex hormone receptors are expressed in different brain areas, such as the hippocampus, hypothalamus, cortex, cerebellum, medial amygdala, substantia nigra, and ventral tegmental area 13 — The active receptor binds to specific DNA sequences named hormone response elements HREs located within the regulatory regions of target genes. The receptor also recruits coactivators and chromatin remodeling complexes that enhance the attachment of the basal transcription machinery to induce gene expression.
Genes that lack HRE can be hormonally induced through the interaction of the receptor with transcription factors like Sp1 and Ap1 19 — Once the receptor dissociates from the DNA, it is marked for degradation through the 26S proteasome 23 These mechanisms are regulated through PR and ER located in the cytoplasm, nucleus, or plasma membrane 30 — 32 or through other membrane receptors that have different biochemical and pharmacological properties 33 These signaling pathways may eventually induce gene transcription. The different mechanisms of action of sex hormones may account for the diverse signaling profiles observed in various brain regions.
The effects of E2 and P4 in the brain depend on hormonal levels and receptor expression. The levels of P4 and E2 fluctuate throughout the life span of the rat modifying different parts of the CNS and causing diverse alterations in brain anatomy, physiology, and behavior 35 E2- and P4-induced plasticity occurs when neuronal cells dynamically respond to hormonal stimuli by modifying its connectivity network and biochemical composition. Brain plasticity can be long lasting, and even the same stimuli can induce different plastic responses at different ages The most dramatic change induced by sex hormones in brain is the driving of its sexual differentiation.
During the fetal—neonatal period, sex hormones permanently modify the brain architecture 13 Neurogenesis, cell differentiation, synaptogenesis, axon guidance, myelination, cell migration, and cell death are some of the main mechanisms occurring during sexual differentiation of the brain. These mechanisms alter the brain area, volume, cell number, cytoarchitecture, cell activity, synaptic connectivity, and neurochemical content 1 After brain differentiation, sex hormone levels in the brain are transitory and fluctuating, and induce the continuous functional adaptation of the CNS throughout the life span of the animal, particularly in females 35 The main periods where sex hormone levels fluctuate during the life span are the beginning of puberty, reproductive cycles, pregnancy, and menopause.
During these phases, alterations in the number of neurons and synapses, glial complexity, morphological variations in dendrites and synapses, and changes in neurotransmitter levels have been reported 41 — These changes promote neuronal and glial remodeling that is critical for cognition, learning, and memory.
For example, spatial working memory varies during se pregnancy, and the memory retention enhanced by E2 wex maintained by P4 45 Further data show that E2 and P4 modify neuronal morphology of the hippocampus of rats and sxe, an important region for memory consolidation 47 It has been recently reported that Pkna enhances object recognition memory consolidation through mTOR and Wnt signaling There is also evidence that both E2 and P4 can modulate GABAergic, dopaminergic, glutamatergic, and serotoninergic neurotransmission, as well as the release of a variety of growth factors from the astroglia 50 — Selection of study sites with adequate representation of women; 3.
The largest cardiopulmonary exercise testing data ever acquired in women at baseline and during trial conduct, and 4. An aggressive approach to optimize background medical therapy that should minimize the differences between treatment groups. The six minute walk test was performed at baseline as well according to standard protocol and patients were asked to complete the Kansas City Cardiomyopathy Questionnaire 21 and Beck Depression Inventory II. Baseline characteristics were compared between men and women using the chi square test for categorical, and t test for continuous variables.
Both the homo VO2 and the 6 min walk homo were significantly lower in the women than in the men. Actin filaments, microtubules MTsand homo filaments, as well as the proteins associated with them, play a homo homo in homo and dendritic homo homo 2.
All tests were performed at the 0. Statistically significant interactions were represented ssex by plotting the predicted CPX outcomes corresponding to a range of values for the significant interaction terms among men and women. The baseline demographics and laboratories of the population by sex are depicted in Table 1. On average, the women were younger than the men and a larger proportion of the African Americans were women. The average EF was higher in the women. The women were more likely to have HF from a non-ischemic etiology and less likely to have a history of hypertension or diabetes. Women were less likely to smoke at any time. However, both peak VO2 and 6 min walk were statistically significantly lower in the women when compared to the men.
Table 1 Subject baseline characteristics and laboratories by sex: