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Article

Daniel J. Bernard, Yining Li, Chirine Toufaily, and Gauthier Schang

The gonadotropins, follicle-stimulating hormone (FSH) and luteinizing hormone (LH), are glycoproteins produced by gonadotrope cells of the anterior pituitary gland. The two hormones act on somatic cells of the gonads in both males and females to regulate fundamental aspects of reproductive physiology, including gametogenesis and steroidogenesis. In males, LH stimulates testosterone production and sperm maturation. FSH also regulates spermatogenesis, though the importance of the hormone in this process differs across species. In females, FSH stimulates ovarian follicle maturation. Follicles are structures composed of oocytes surrounded by two types of somatic cells, granulosa and theca cells. FSH stimulates granulosa cells to proliferate and to increase their production of the aromatase enzyme. LH stimulates theca cells to make androgens, which are converted into estrogens by aromatase in granulosa cells. A surge of LH also stimulates ovulation of mature follicles. Gonadotropin-releasing hormone (GnRH) from the brain is the principal stimulator of gonadotropin synthesis and secretion from the pituitary. The sex steroids (androgens and estrogens) that are produced by the gonads in response to the gonadotropins feedback to the brain and pituitary gland. In the brain, these hormones usually slow the release of GnRH through a process called negative feedback, which in turn leads to decreases in FSH and LH. The steroids also modulate the sensitivity of the pituitary to GnRH in addition to directly regulating expression of the genes that encode the gonadotropin subunits. These effects are gene- and species-specific. In females, estrogens also have positive feedback actions in the brain and pituitary in a reproductive cycle stage-dependent manner. This positive feedback promotes GnRH and LH release, leading to the surge of LH that triggers ovulation. The gonadotropins are dimeric proteins. FSH and LH share a common α-subunit but have hormone-specific subunits, FSHβ and LHβ. The β subunits provide a means for differential regulation and action of the two hormones. In the case of FSH, there is a second gonadal feedback system that specifically regulates the FSHβ subunit. The gonads produce proteins in the transforming growth factor β (TGFβ) family called inhibins, which come in two forms (inhibin A and inhibin B). The ovary produces both inhibins whereas the testes make inhibin B alone. Inhibins selectively suppress FSH synthesis and secretion, without affecting LH. The pituitary produces additional TGFβ proteins called activins, which are structurally related to inhibins. Activins, however, stimulate FSH synthesis by promoting transcription of the FSHβ subunit gene. Inhibins act as competitive receptor antagonists, binding to activin receptors and blocking activin action, and thereby leading to decreases in FSH. Together, GnRH, sex steroids, activins, and inhibins modulate and coordinate gonadotropin production and action to promote proper gonadal function and fertility.

Article

Paul E. Micevych and Melinda A. Mittelman-Smith

In the last two decades of the 20th century, key findings in the field of estrogen signaling completely changed our understanding of hormones: first, steroidogenesis was demonstrated in the CNS; second, a vast majority of cells in the nervous system were shown to have estrogen receptors; third, a second nuclear estrogen receptor (ERß) was cloned; and finally, “nuclear” receptors were shown to be present and functional in the cell membrane. Shortly thereafter, even more membrane estrogen receptors were discovered. Steroids (estrogens, in particular) began to be considered as neurotransmitters and their receptors were tethered to G protein-coupled receptor signaling cascades. In some parts of the brain, levels of steroids appeared to be independent of those found in the circulation and yet, circulating steroids had profound actions on the brain physiology. In this review, we discuss the interaction of peripheral and central estrogen action in the context of female reproduction—one of the best-studied aspects of steroid action. In addition to reviewing the evidence for steroidogenesis in the hypothalamus, we review membrane-localized nuclear receptors coupling to G protein-signaling cascades and the downstream physiological consequences for reproduction. We will also introduce newer work that demonstrates cell signaling for a common splice variant of estrogen receptor-α (ERα), and membrane action of neuroprogesterone in regulating estrogen positive feedback.

Article

Anthony C. Hackney and Eser Ağgön

Stress is encountered by every individual on a daily basis. Such encounters can be of a negative (distress) or a positive (eustress) nature. Excessive and chronic distress exposure is associated with numerous health problems affecting both physiological and psychological components of a person’s well-being. One mediating aspect of these occurrences is the responses of the neuroendocrine system with the body. Physical activity (i.e., exercise) produces large and dramatic changes in the neuroendocrine system as it serves as a “stressor” to the system. To this end, though, chronic engagement in physical activity leads to exercise training-induced adaptations within the neuroendocrine system that potentiate an individual’s ability to deal with distressful experiences and exposures. Therefore, becoming more physically fit and exercise trained is one potential adjunctive therapy available for clinicians to recommend in the treatment of health problems associated with chronic exposure to distress.

Article

Ashlyn Swift-Gallant and S. Marc Breedlove

While prenatal sex hormones guide the development of sex-typical reproductive structures, they also act on the developing brain, resulting in sex differences in brain and behavior in animal models. Stemming from this literature is the prominent hypothesis that prenatal neuroendocrine factors underlie sex differences in human sexual orientation, to explain why most males have a preference for female sexual partners (gynephilia), whereas most females display a preference for male sexual partners (androphilia). Convergent evidence from experiments of nature and indirect markers of prenatal hormones strongly support a role for prenatal androgens in same-same sexual orientations in women, although this finding is specific to a subset of lesbians who are also gender nonconforming (“butch”). More gender-conforming lesbians (“femmes”) do not show evidence of increased prenatal androgens. The literature has been more mixed for male sexual orientation: some report evidence of low prenatal androgen exposure, while others report evidence of high androgen levels and many other studies find no support for a role of prenatal androgen exposure in the development of androphilia in males. Recent evidence suggests there may be subgroups of gay men who owe their sexual orientation to distinct biodevelopmental mechanisms, which could account for these mixed findings. Although this research is young, it is similar to findings from lesbian populations, because gay men who are more gender nonconforming, and report a preference for receptive anal sex, differ on markers of prenatal development from gay men who are more gender conforming and report a preference for insertive anal sex. This chapter concludes with future research avenues including assessing whether multiple biodevelopmental pathways underlie sexual orientation and whether neuroendocrine factors and other biological mechanisms (e.g., immunology, genetics) interact to promote a same-sex sexual orientation.

Article

Dayna L. Averitt, Rebecca S. Hornung, and Anne Z. Murphy

The modulatory influence of sex hormones on acute pain, chronic pain disorders, and pain management has been reported for over seven decades. The effect of hormones on pain is clearly evidenced by the multitude of chronic pain disorders that are more common in women, such as headache and migraine, temporomandibular joint disorder, irritable bowel syndrome, chronic pelvic pain, fibromyalgia, rheumatoid arthritis, and osteoarthritis. Several of these pain disorders also fluctuate in pain intensity over the menstrual cycle, including headache and migraine and temporomandibular joint disorder. The sex steroid hormones (estrogen, progesterone, and testosterone) as well as some peptide hormones (prolactin, oxytocin, and vasopressin) have been linked to pain by both clinical and preclinical research. Progesterone and testosterone are widely accepted as having protective effects against pain, while the literature on estrogen reports both exacerbation and attenuation of pain. Prolactin is reported to trigger pain, while oxytocin and vasopressin have analgesic properties in both sexes. Only in the last two decades have neuroscientists begun to unravel the complex anatomical and molecular mechanisms underlying the direct effects of sex hormones and mechanisms have been reported in both the central and peripheral nervous systems. Mechanisms include directly or indirectly targeting receptors and ion channels on sensory neurons, activating pain excitatory or pain inhibitory centers in the brain, and reducing inflammatory mediators. Despite recent progress, there remains significant controversy and challenges in the field and the seemingly pleiotropic role estrogen plays on pain remains ambiguous. Current knowledge of the effects of sex hormones on pain has led to the burgeoning of gender-based medicine, and gaining further insight will lead to much needed improvement in pain management in women.

Article

Lily Yan, Laura Smale, and Antonio A. Nunez

Circadian rhythms are endogenous daily rhythms evident in behavior and physiology. In mammals, these rhythms are controlled by a hierarchical network of oscillators showing a coherent circadian coordination or coupling. The hypothalamic suprachiasmatic nucleus (SCN) sits on top of the hierarchy and coordinates the phase of oscillators in other brain regions and in peripheral organs, including endocrine glands. The phase of the SCN oscillator, in reference to the daily light-dark cycle, is identical across mammalian species regardless of whether they are most active during the day or night, that is, diurnal or nocturnal. However, the extra-SCN or peripheral oscillators are out of phase and are often reversed by 180° across diurnal and nocturnal mammals. In the endocrine system, with the notable exception of the pattern of pineal melatonin secretion, which features elevated levels at night regardless of the activity profile of the species, most endocrine rhythms show a 180° reversal when diurnal and nocturnal species are compared. There is also evidence of differences between nocturnal and diurnal species with respect to their rhythms in sensitivity or responsiveness to hormonal stimulation. One of the major unanswered questions in the field of comparative endocrinology relates to the mechanism responsible for the differential coupling in diurnal and nocturnal mammals of extra-SCN oscillators and overt circadian rhythms with the SCN oscillator and the light dark cycle. Viable hypotheses include species-specific switches from excitation to inhibition at key nodes between the SCN and its targets, the presence of extra-SCN signals that converge on SCN targets and reverse the outcome of SCN signals, and changes in oscillatory parameters between the oscillator of the SCN and those outside the SCN resulting in an anti-phase coupling among key oscillators.

Article

Kory Floyd and Colter D. Ray

Affectionate communication comprises the verbal and nonverbal behaviors people use to express messages of love, appreciation, fondness, and commitment to others in close relationships. Like all interpersonal behaviors, affectionate communication has biological and physiological antecedents, consequences, and correlates, many of which have implications for physical health and wellness. Investigating these factors within a biological framework allows for the adjudication of influences beyond those attributable to the environment. In particular, there are observable genetic and neurological differences between individuals with a highly affectionate disposition and those less prone to communicating affection, suggesting that variance in the tendency to engage in affectionate behavior is not entirely the result of environmental influences such as enculturation, parenting, and media exposure. In addition, the expression of affection is associated with markers of immune system competence and appears to help the body to relax and remain calm. The biological effects of affectionate communication are perhaps most pronounced in situations involving either acute or chronic stress. Specifically, highly affectionate individuals are less likely than others to overreact physiologically to stress-inducing events. Whatever stress reaction they do mount is better regulated than among their less affectionate counterparts. Moreover, highly affectionate individuals—or simply those who receive expressions of affection prior to or immediately following a stressful situation—exhibit faster physiological recovery from their elevated stress. Perhaps unsurprisingly, therefore, being deprived of adequate affectionate communication is predictive of multiple physical and psychological detriments, including elevated stress and exacerbated depression, social and relational problems, insecure attachment, susceptibility to diagnosed anxiety and mood disorders, susceptibility to diagnosed secondary immune disorders, chronic pain, and sleep disturbances.

Article

Understanding of the brain mechanisms regulating reproductive behaviors in female laboratory animals has been aided greatly by our knowledge of estrogen receptors in the brain. Hypothalamic neurons that express the gene for estrogen receptor-alpha regulate activity in the neural circuit for the simplest female reproductive response, lordosis behavior. In turn, many of the neurotransmitter inputs to the critical hypothalamic neurons have been studied using electrophysiological and neurochemical techniques. The upshot of all of these studies is that lordosis behavior presents the best understood set of mechanisms for any mammalian behavior.

Article

Aleksander Ksiazkiewicz and Seyoung Jung

The study of biology and politics is rapidly moving from being an isolated curiosity to being an integral part of the theories that political scientists propose. The necessity of adopting this interdisciplinary research philosophy will be increasingly apparent as political scientists seek to understand the precise mechanisms by which political decisions are made. To demonstrate this potential, scholars of biopolitics have addressed common misconceptions about biopolitics research (i.e., the nature-nurture dichotomy and biological determinism) and used different methods to shed light on political decision making since the turn of the 21st century—including methods drawn from evolutionary psychology, genomics, neuroscience, psychophysiology, and endocrinology. The field has already come far in its understanding of the biology of political decision making, and several key findings have emerged in biopolitical studies of political belief systems, attitudes, and behaviors. This area of research sheds light on the proximate and ultimate causes of political cognition and elucidates some of the ways in which human biology shapes both the human universals that make politics possible and the human diversity that provides it with such dynamism. Furthermore, three emerging areas of biopolitics research that anticipate the promise of a biologically informed political science are research into gene-environment interplay, research into the political causes and consequences of variation in human microbiomes, and research that integrates chronobiology—the study of the biological rhythms that regulate many aspects of life, including sleep—into the study of political decision making.

Article

Eliot A. Brenowitz

Animals produce communication signals to attract mates and deter rivals during their breeding season. The coincidence in timing results from the modulation of signaling behavior and neural activity by sex steroid hormones associated with reproduction. Adrenal steroids can influence signaling for aggressive interactions outside the breeding season. Androgenic and estrogenic hormones act on brain circuits that regulate the motivation to produce and respond to signals, the motor production of signals, and the sensory perception of signals. Signal perception, in turn, can stimulate gonadal development.

Article

Vermiculture is the art, science, and industry of raising earthworms for baits, feeds, and composting of organic wastes. Composting through the action of earthworms and microogranisms is commonly referred to as vermicomposting. Vermiculture is an art because the technology of raising earthworms requires a comprehensive understanding of the basic requirements for growing earthworms in order to design the space and the system by which organic wastes can be processed efficiently and successfully. It is a science because the technology requires a critical understanding and consideration of the climatic requirements, nutritional needs, growth cycles, taxonomy, and species of earthworms suitable for vermicomposting in order to develop a working system that supports earthworm populations to process successfully the intended organic wastes. The nature of the organic wastes also needs to be taken into careful consideration, especially its composition, size, moisture content, and nutritional value, which will eventually determine the overall quality of the vermicomposts produced. The quality of organic wastes also determines the ability of the earthworms to consume and process them, and the rate by which they turn these wastes into valuable organic amendments. The science of vermiculture extends beyond raising earthworms. There are several lines of evidence that vermicomposts affect plant growth significantly. Vermiculture is an industry because it has evolved from a basic household bin technology to commercially scaled systems in which economic activities emanate from the cost and value of obtaining raw materials, the building of systems, and the utilization and marketing of the products, be they in solid or aqueous extract forms. Economic returns are carefully valued from the production phase to its final utilization as an organic amendment for crops. The discussion revolves around the development of vermiculture as an art, a science, and an industry. It traces the early development of vermicomposting, which was used to manage organic wastes that were considered environmentally hazardous when disposed of improperly. It also presents the vermicomposting process, including its basic requirements, technology involved, and product characteristics, both in solid form and as a liquid extract. Research reports from different sources on the performance of the products are also provided. The discussion attempts to elucidate the mechanisms involved in plant growth and yield promotion and the suppression of pests and diseases. Certain limitations and challenges that the technology faces are presented as well.

Article

Conscience P. Bwiza, Jyung Mean Son, and Changhan Lee

Aging is a progressive process with multiple biological processes collectively deteriorating with time, ultimately causing loss of physiological functions necessary for survival and reproduction. It is also thought to have a strong evolutionary basis, largely resulting from the lack of selection force. Here, we discuss the evolutionary aspects of aging and a selection of theories founded on a variety of biological functions that have been shown to be involved in aging in multiple model organisms, ranging from the simple yeast, worms, flies, killifish, and rodents, to non-human primates and humans. The conglomerate of distinct theories has together revolutionized aging research in the past several decades, far more than what humankind has known since the dawn of civilization. However, not one theory alone can independently explain aging and should not be interpreted out of context of the cell and organism in its entirety. That said, the 21st century has been and will be an exciting time in the field of aging, with scientific advances on health span and lifespan being made at multiple fronts of biology and medicine in an unprecedented scale.

Article

Vanessa L. Burrows

Stress has not always been accepted as a legitimate medical condition. The biomedical concept stress grew from tangled roots of varied psychosomatic theories of health that examined (a) the relationship between the mind and the body, (b) the relationship between an individual and his or her environment, (c) the capacity for human adaptation, and (d) biochemical mechanisms of self-preservation, and how these functions are altered during acute shock or chronic exposure to harmful agents. From disparate 19th-century origins in the fields of neurology, psychiatry, and evolutionary biology, a biological disease model of stress was originally conceived in the mid-1930s by Canadian endocrinologist Hans Selye, who correlated adrenocortical functions with the regulation of chronic disease. At the same time, the mid-20th-century epidemiological transition signaled the emergence of a pluricausal perspective of degenerative, chronic diseases such as cancer, heart disease, and arthritis that were not produced not by a specific etiological agent, but by a complex combination of multiple factors which contributed to a process of maladaptation that occurred over time due to the conditioning influence of multiple risk factors. The mass awareness of the therapeutic impact of adrenocortical hormones in the treatment of these prevalent diseases offered greater cultural currency to the biological disease model of stress. By the end of the Second World War, military neuropsychiatric research on combat fatigue promoted cultural acceptance of a dynamic and universal concept of mental illness that normalized the phenomenon of mental stress. This cultural shift encouraged the medicalization of anxiety which stimulated the emergence of a market for anxiolytic drugs in the 1950s and helped to link psychological and physiological health. By the 1960s, a growing psychosomatic paradigm of stress focused on behavioral interventions and encouraged the belief that individuals could control their own health through responsible decision-making. The implication that mental power can affect one’s physical health reinforced the psycho-socio-biological ambiguity that has been an enduring legacy of stress ever since. This article examines the medicalization of stress—that is, the historical process by which stress became medically defined. It spans from the mid-19th century to the mid-20th century, focusing on these nine distinct phases: 1. 19th-century psychosomatic antecedent disease concepts 2. The emergence of shell-shock as a medical diagnosis during World War I 3. Hans Selye’s theorization of the General Adapation Syndrome in the 1930s 4. neuropsychiatric research on combat stress during World War II 5. contemporaneous military research on stress hormones during World War II 6. the emergence of a risk factor model of disease in the post-World War II era 7. the development of a professional cadre of stress researchers in the 1940s and 50s 8. the medicalization of anxiety in the early post–World War II era 9. The popularization of stress in the 1950s and pharmaceutical treatments for stress, marked by the cultural assimilation of paradigmatic stress behaviors and deterrence strategies, as well pharmaceutical treatments for stress.