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Corresponding Author Peter B. Gray, Associate Professor. In this mini-review, we consider an evolutionary biocultural perspective on human aging and sexuality. An evolutionary approach to senescence highlights the energetic trade-offs between fertility and mortality. By comparing humans to other primates, we situate human senescence as an evolutionary process, with shifts in postreproductive sexual behavior in this light. Age-related declines in sexual behavior are typical for humans but also highly contingent on the sociocultural context within which aging individuals express their sexuality.

We briefly review some of the most comprehensive studies of aging and sexual behavior, both from the USA and cross-culturally. We frame these patterns with respect to the long-term relationships within which human sexual behavior typically occurs. Because sexuality is typically expressed within pair-bonds, sexual behavior sometimes declines in both members of a couple with age, but also exhibits sex-specific effects that have their roots in evolved sex differences.

In this mini-review, we consider an evolutionary biocultural perspective on aging and sexuality, highlighting some of the most comprehensive studies of aging and sexual behavior, both from the USA and internationally. We find that age-related declines in human sexual behavior are typical but also highly contingent on the sociocultural context within which aging individuals express their sexuality, as the above quote describing the Aka illustrates.

Because both men and women typically express their sexuality within long-term relationships, their sexual behavior sometimes changes in tandem i. An evolutionary perspective on aging recognizes that species should invest in maintenance functions such as immune systems in mammals if age-specific fertility and mortality pay to do so. If, for example, members of a species continue to reproduce and are subject to sufficiently low extrinsic mortality mortality that is difficult to avoid, such as predationthen they can afford to maintain themselves, investing in their own capacities for continued survival, and in turn living long lives.

There is empirical cross-species support for this body of theory. To illustrate, let us contrast mainland opossums living in the south of the USA with opossums living on a nearby island, where they had been isolated from predators for thousands of years. Austad [ 2 ] predicted and found support for the hypothesis that the island opossums would senesce at Ladies seeking sex Onward Indiana rates than their mainland cousins. Under the idea that islanders faced lower extrinsic mortality due to lower rates of predation, he found that island opossums died at slower rates with advancing age meaning the average islander lived a longer life and also scored lower on a physiological measure of stress wear and tear to their tail tendon fibers.

As other examples, bats tend to outlive their similarly sized mammalian cousins such as squirrels, a pattern consistent with theory since bats can more readily escape would-be predators by flying away, enabling them to invest in maintenance functions [ 2 ]. Tortoises, like those Charles Darwin encountered in the Galapagos Islands, tend to lead long lives, in part because they hold protection hard shells literally on their backs. Such examples illustrate how fertility and mortality schedules shape the rate of senescence loss of function with age among organisms.

The disposable soma theory articulates an evolutionary perspective on aging, by considering the life history trade-offs involved with maintenance. These trade-offs ultimately impact reproductive function, not as a result of population control as some have arguedbut as a result of natural energetic concessions [ 3 ].

Once an organism reaches an age when survivorship in the wild is highly unlikely, the state of its somatic integrity, or otherwise, is inconsequential. What light does this evolutionary perspective shed on human aging? Humans senesce at slower rates than our great ape cousins, and apes, in turn, senesce at slower rates compared with other primates such as lemurs and both old-world monkeys e.

As an example, wild chimpanzee mortality data from five field sites in Africa showed that almost every chimpanzee was dead by age 50 [ 4 ]. By contrast, recent studies of human hunter-gatherers and horticulturalists such as the! Kung of the Kalahari, the Hadza of Tanzania, the Ache of Paraguay and the Tsimane of Bolivia have found that these human populations have longer lives than our ape cousins living either in captivity or the wild [ 5 ].

Of course, recent human hunter-gatherers also have technological and cognitive shields controlled use of fire, projectile weapons, knowledgeable elders against Ladies seeking sex Onward Indiana insults that our more distant foraging ancestors lacked.

How far back, then, we can project mortality patterns from recent foragers to more distant foragers is an open question. The available bioarchaeological samples e. When combining this line of evidence with other paleoanthropological data on rates of development, brain size estimates and archaeological evidence of tool use and subsistence, along with the comparative ape and contemporary hunter-gatherer data, the best guess would be that the slower rate of human senescence evolved in the genus Homo at some point within the past 2 million years [ 6 ].

A more specific educated guess would be that only with modern humans ourselves within the pastor so years have people been living long enough to even contemplate a postreproductive existence [ 7 ].

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Considerable debate has arisen concerning why human females, in particular, live well beyond their reproductive capacities [ 8910 ]. An extended life span gives rise to postreproductive female existence. While some argue that the cause of extended human life spans is grandmaternal care i. Several arguments favor the latter byproduct view, focusing on the fact that selection tends to be relatively weak or absent for any traits specific to postreproductive years of life.

The comparative nonhuman evidence convincingly demonstrates that postreproductive female lives are hardly unique to humans. However, even among forager groups, humans appear to spend a ificantly longer proportion of their lives in postreproductive states than is the case in postreproductive captive animals.

Interestingly, peri- and postmenopausal gorillas no longer maintained the same social affiliations with a silverback adult male gorillathough whether that is because females were not interested in doing so or males were not as interested in the females, or both, we do not know [ 13 ].

Put another way, the overall theoretical and empirical evidence is most consistent with the view that human senescence has a of parallels with captive nonhuman animals and has slowed recently in hominin evolution perhaps only with modern humans ourselves. This has given rise to Ladies seeking sex Onward Indiana and extended postreproductive lives that can be spent having sex, among other things.

While most of the focus on human reproductive aging has centered on females, male senescence can also be understood in an evolutionary perspective [ 1516 ]. Estimates of age-specific fertility in forager societies such as the! The fact that human sexual and reproductive behavior typically occurs within long-term slightly polygynous bonds is noteworthy, as this social context differs from our great ape cousins and was likely derived within the past 2 million years of Homo evolution [ 1618 ]. Yet, male reproductive potential does decline with age, with age-specific fertility quite low by the 50s and onwards.

In a prospective fecundability study of couples women aged 18—40 yearsboth men and women experienced increased infertility with age [ 19 ]. Humans, like most other species of mammals, also exhibit sex differences in senescence, which ultimately can be traced to sexual selection.

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Across various studies of mammals and birds in the wild, polygynous species typically display sex differences in mortality males having shorter lives, on averagewhereas among monogamous species, male and female mortality profiles are similar [ 20 ].

Human males tend to live shorter lives on average than females, although at lessened intensity compared with, say, orangutans or gorillas. An evolutionary perspective on senescence helps make sense of age-related changes in the proximate mechanisms regulating sexual response [ 2122 ]. For both females and males, general frailty declines at slower rates than our closest primate cousins, making it easier for older humans to remain sexually active they are alive, and better able to function, though subject to the influence of declining health.

Female peak fecundity occurs at ages 25—35, declines toward menopause and ceases altogether at menopause. With advancing age, the ovary reduces its capacity to both produce a fertilizable egg and release sex steroids that facilitate sexual response.

Age-related declines in estrogen, especially during the menopausal transition, negatively impact vaginal lubrication, elasticity and vasocongestion and can also reduce sexual desire. Male reproductive function reveals declines in testosterone by the early 30s onwards, although there is also population variation in the pattern of decline. Male declines in sexual desire and erectile function are more pronounced from the 60s onwards and may, in part, be traced to age-related declines in testosterone. All said, the mechanisms of female and male sexual function decline with age, Ladies seeking sex Onward Indiana that can be understood with respect to the evolutionary background highlighted above and that would anticipate age-related changes in actual sexual behavior.

These age-related changes in reproductive physiology also point to age-related shifts in aggression, courtship and childcare e. In parts of India, a couple is supposed to stop engaging in sex once their eldest son marries, a sexual recognition of shifting family priorities. In the 19th century Oneida commune of upstate New York, young men were sexually kept by postmenopausal women.

There seems to be less danger attached to it, since husbands are often away for long periods or perhaps are less jealous.

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In light of these anecdotes, what about broader patterns of sexual behavior associated with human aging? Here, we present the of several large, recent quantitative US studies, before also considering patterns drawn from several international studies. Cropped and reprinted, with permission, from David Steinberg. In the USA, the frequency of sexual intercourse over a 1-year period was found to be inversely related to age, with Ladies seeking sex Onward Indiana 18—30 years reportedly averaging sex 78 times per year, compared to those 60—69 years averaging 23 times per year [ 24 ].

More recently, inLindau et al. By asking whether an individual had sex in the 12 months, it was found that the proportion of men and women sexually active declined with age but was lower at all ages among women. Thus, the falloff in sex with advancing age is a robust feature. As for sexual practices, the vast majority entails vaginal intercourse. Of other sexual practices, rates of both oral sex and masturbation drop off steeply with advancing age. The drop in masturbation, which does not require the same negotiations with a partner as oral sex or intercourse, suggests that other factors such as declining sexual desire are wielding measurable effects on age-related changes in sexual behavior.

It is worth noting that fairly recent historical shifts in sexual culture, including the hierarchical reordering of oral sex and intercourse [ 26 ], may also result in a cohort effect with respect to rates of specific sexual behavior. How should we interpret the sex differences in age-related patterns of sexuality?

One view is that these may be indicative of sex differences in sexual desire that persist at advancing ages; that interpretation could be most sensible with respect to sex differences in masturbation. However, another major cause of these patterns is that the demographic landscape changes with advancing age; with men often older than their sexual partners, men dying at younger ages and women less likely to remarry, the demographics at older ages increasingly tilt toward living females. Since most sex occurs within long-term relationships such as marriage, older women simply have fewer sexual options available to them.

Poor self-reported health was also associated with lower rates of sexual behavior, in females and males alike. In addition to age-related health problems that may impact sexual activity, a variety of medications taken for age-related conditions have side effects of reducing sexual desire and performance. An even more recent US survey in National Survey of Sexual Health and Behavior yielded generally similar patterns of age-related changes in sexual behavior, although relying on a wider age spread participants 14—94 years of age and different methods e.

The age-related declines and lower rates of sexual behaviors among females appeared in this study as in the earlier one.

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Also of note, among men aged 50—59 years and older, a higher percentage reported performing and receiving oral sex from another male compared with the rates of females performing and receiving oral sex from another female, an indication that in advanced ages the rates of same-sex male sexual behavior were higher than the rates of same-sex female sexual behavior. One of the best studied aspects of aging and human sexuality has been across the menopausal transition.

While aging itself may influence sexuality, a related question has concerned effects specifically traced to the physiological effects of menopause itself. Reviewing approximately 15 studies, with both cross-sectional and longitudinal des, Dennerstein et al. Providing an international view of aging and sexuality, Laumann et al. The data were presented differently, not allowing direct comparisons with the US data by age or Ladies seeking sex Onward Indiana with regard to rates of sexual behavior, as the focus was on clustering the global patterns of aging and sexuality by attitudinal and relationship factors.

As a central finding, there was a positive relationship across countries in the percentage of participants who reported satisfaction with their sexual function and relationship satisfaction. Important messages, then, from this international study are that sexual satisfaction and relationship satisfaction are highly related no surprise given the primacy of long-term relationships in human sexuality and that considerable variation in sexual and relationship satisfaction exists among aged populations.

Another important finding from Ladies seeking sex Onward Indiana international study [ 29 ] concerns sex differences in reported sexual well-being. Men reported higher levels of physical pleasure, emotional pleasure, satisfaction with sexual function and importance of sex than did women.

In other words, sex in companionate relationships serves not only reproductive purposes, but also expresses the quality of the relationship. They found that older males were sexually active in 20 of 28 societies, and older females were sexually active in 22 of 26 societies from which they could obtain data. Thus, the admittedly slim Human Relations Area Files data indicate that older women and men are quite commonly sexually active with advancing age. One means by which older men remain sexually active in some societies is by marrying younger wives, sometimes in polygynous unions.

The desire to have more children is sometimes articulated as a reason for older men remaining sexually active with their younger wives, as among Kuwaiti Bedouin and the Chukchee. However, a refrain in at least five societies, including among prospective young brides in the Trobriands and among the Lovedu of South Africa, is that old men are not physically attractive.

In some other cases, such as the Taiwan Hokkien, the thought that older men and women remain sexually active is found distasteful, but for a different reason, namely that they already have grandchildren. Here, it is expected that grandparents will sacrifice their sex lives to focus on raising their grandchildren. One of the noteworthy patterns of female sexuality is that reports from 18 of 22 societies indicated that women had sex with younger men, sometimes considerably younger.

In some cases, it appears that their younger male partners were desirable, with some of them married men. However, in the majority of cases this seems to reflect a context in which older women are not viewed as most desirable, but rather taken as partners because they are the only ones available to some men.

Older women were also utilized by young boys as sexual initiators and sexual instructors in several of the societies reported, as in the Tupinamba of northeast Brazil Given marital customs whereby a young husband might marry a much older woman e. Another noteworthy pattern in female sexuality was that older women frequently experienced a loosening of constraints to their sexual expression.

With their reproductive capacities past, older women might make ribald jokes and display an enhanced sexual aggressiveness, as among the Taiwanese Hokkien and the Muria of India.

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