Extrinsic Mortality Risk Research Articles

Mortality risk predicts global, local, and individual patterns of human reproduction

BackgroundHuman reproductive dynamics in the post-industrial world are typically explained by economic, technological, and social factors including the prevalence of contraception and increasing numbers of women in higher education and the workforce. These factors have been targeted by multiple world governments as part of family policies, yet those policies have had limited success. The current work adopts a life history perspective from evolutionary biology: like most species, human populations may respond to safer environments marked by lower morbidity and mortality by slowing their reproduction and reducing their number of offspring. We test this association on three levels of analysis using global, local, and individual data from publicly available databases.ResultsData from over 200 world nations, 3,000 U.S. counties and 2,800 individuals confirm an association between human reproductive outcomes and local mortality risk. Lower local mortality risk predicts “slower” reproduction in humans (lower adolescent fertility, lower total fertility rates, later age of childbearing) on all levels of analyses, even while controlling for socioeconomic variables (female employment, education, contraception).ConclusionsThe association between extrinsic mortality risk and reproductive outcomes, suggested by life history theory and previously supported by both animal and human data, is now supported by novel evidence in humans. Social and health policies governing human reproduction, whether they seek to boost or constrain fertility, may benefit from incorporating a focus on mortality risk.

Attitudes and Laws About Abortion Are Linked to Extrinsic Mortality Risk: A Life-History Perspective on Variability in Reproductive Rights

Abortion policy is conventionally viewed as a political matter with religious overtones. This article offers a different view. From the perspective of evolutionary biology, abortion at a young age can represent prioritization of long-term development over immediate reproduction, a pattern established in other animal species as resulting from stable ecologies with low mortality risk. We examine whether laws and moral beliefs about abortions are linked to local mortality rates. Data from 50 U.S. states, 202 world societies, 2,596 adult individuals in 363 U.S. counties, and 147,260 respondents across the globe suggest that lower levels of mortality risk are associated with more permissive laws and attitudes toward abortion. Those associations were observed when we controlled for religiosity, political ideology, wealth, education, and industrialization. Integrating evolutionary and cultural perspectives offers an explanation as to why moral beliefs and legal norms about reproduction may be sensitive to levels of ecological adversity.

Comparative life history patterns of female gorillas.

Several theories have been proposed to explain the impact of ecological conditions on differences in life history variables within and between species. Here we compare female life history parameters of one western lowland gorilla population (Gorilla gorilla gorilla) and two mountain gorilla populations (Gorilla beringei beringei). We compared the age of natal dispersal, age of first birth, interbirth interval, and birth rates using long-term demographic datasets from Mbeli Bai (western gorillas), Bwindi Impenetrable National Park and the Virunga Massif (mountain gorillas). The Mbeli western gorillas had the latest age at first birth, longest interbirth interval, and slowest surviving birth rate compared to the Virunga mountain gorillas. Bwindi mountain gorillas were intermediate in their life history patterns. These patterns are consistent with differences in feeding ecology across sites. However, it is not possible to determine the evolutionary mechanisms responsible for these differences, whether a consequence of genetic adaptation to fluctuating food supplies ("ecological risk aversion hypothesis") or phenotypic plasticity in response to the abundance of food ("energy balance hypothesis"). Our results do not seem consistent with the extrinsic mortality risks at each site, but current conditions for mountain gorillas are unlikely to match their evolutionary history. Not all traits fell along the expected fast-slow continuum, which illustrates that they can vary independently from each other ("modularity model"). Thus, the life history traits of each gorilla population may reflect a complex interplay of multiple ecological influences that are operating through both genetic adaptations and phenotypic plasticity.

Being Cared for and Growing Up Slowly: Parenting Slows Human Life History

SYNOPSIS Objective. For most animals, extrinsic mortality risks drive a fast life history (LH) strategy in which animals disregard risks and accelerate reproduction. Instead of perpetuating mortality driving fast LH, humans have reduced almost all mortality risks in living environments, resulting in a significant slowing of LH. Additionally, humans exhibit invested parenting which entails teaching their young survival or mortality reduction skills. Could parenting provide an additional pathway to the development and slowing of human LH? Design. Data reported here come from interviews and questionnaires administered to a community sample of 286 rural Chinese parents and their children when the children were on average 7, 8, and 11 years old. Results. Parental acceptance statistically mediates and moderates the longitudinal association between environmental adversities and children’s LH. Conclusions. Parenting breaks the species-general contingency between mortality conditions and fast offspring LH strategies and provides an additional pathway to the development and slowing of human LH.

Food insecurity, diet and mental distress among resource insecure students during COVID-19.

It is well documented that college student populations are vulnerable to food insecurity and other adverse environmental conditions. Additionally, exposure to environmental adversity can have deleterious, long-term effects on physical and mental health. This study applies evolutionary life history theory to examine the relationship between environmental adversity, mental distress and diet among resource insecure university students during the COVID-19 pandemic. Structured and semi-structured surveys were used to assess perceptions of environmental adversity (including mortality risk, food insecurity and resource availability; and changes in these factors over the course of COVID-19), mental distress, diet and use of campus support services. Participants included 51 college students recruited through an economic crisis center located at a large public university in southern California. Most students were experiencing mental distress and food insecurity, and food insecurity and other components of adversity increased during COVID-19. Food insecurity was significantly associated with both perceived extrinsic mortality risk and mental distress, whereas mental distress was significantly associated with reduced dietary quality and caloric intake. Use of two or more campus support resources and/or living with family or rent free disrupted the associations of food insecurity with extrinsic mortality risk and mental distress. This study contributes to a growing body of applied evolutionary frameworks concerned with the health and wellbeing of economically vulnerable populations. It also provides novel insights informed by life history theory into interventions and recommendations for improving support services for financially insecure college students.

Child attachment in adjusting the species-general contingency between environmental adversities and fast life history strategies.

Extrinsic mortality risks calibrating fast life history (LH) represent a species-general principle that applies to almost all animals including humans. However, empirical research also finds exceptions to the LH principle. The present study proposes a maternal socialization hypothesis, whereby we argue that the more human-relevant attachment system adds to the LH principle by up- and down-regulating environmental harshness and unpredictability and their calibration of LH strategies. Based on a longitudinal sample of 259 rural Chinese adolescents and their primary caregivers, the results support the statistical moderating effect of caregiver-child attachment on the relation between childhood environmental adversities (harshness and unpredictability) and LH strategies. Our theorizing and findings point to an additional mechanism likely involved in the organization and possibly the slowdown of human LH.

Extrinsic and Existential Mortality Risk in Reproductive Decision-Making: Examining the Effects of COVID-19 Experience and Climate Change Beliefs.

While the COVID-19 pandemic has presented an immediate risk to human life around the world, climate change poses an arguably greater—although less immediate—threat to our species’ survival. Within the framework of life-history theory (LHT), this pre-registered study investigated whether extrinsic risk (i.e., external factors that pose a risk to an individual’s life, e.g., COVID-19) and existential risk (i.e., risks with outcomes that threaten the existence of humans as a species, e.g., climate change) had similar or different relationships with reproductive decision-making. A UK representative sample of 325 participants between 18 and 35 years of age was asked to indicate their ideal number of children, ideal age to start having children, and whether their desire for a child had recently changed. Participants were asked about their experiences of COVID-19 and given a series of scales with which to assess their beliefs about climate change. In support of LHT, the study found evidence that knowing people who had been hospitalized with or died of COVID-19 was associated with a greater ideal number of children. Conversely, there was no clear evidence of a relationship between climate change beliefs and reproductive decision-making. The repercussions for understanding how we interpret and respond to different forms of mortality risk are discussed.

Antagonistic pleiotropy and the evolution of extraordinary lifespans in eusocial organisms.

Queens of eusocial species live extraordinarily long compared to their workers. So far, it has been argued that these lifespan divergences are readily explained by the classical evolutionary theory of ageing. As workers predominantly perform risky tasks, such as foraging and nest defense, and queens stay in the well‐protected nests, selection against harmful genetic mutations expressed in old age should be weaker in workers than in queens due to caste differences in extrinsic mortality risk, and thus, lead to the evolution of longer queen and shorter worker lifespans. However, these arguments have not been supported by formal models. Here, we present a model for the evolution of caste‐specific ageing in social insects, based on Williams’ antagonistic pleiotropy theory of ageing. In individual‐based simulations, we assume that mutations with antagonistic fitness effects can act within castes, that is, mutations in early life are accompanied by an antagonistic effect acting in later life, or between castes, where antagonistic effects emerge due to caste antagonism or indirect genetic effects between castes. In monogynous social insect species with sterile workers, large lifespan divergences between castes evolved under all different scenarios of antagonistic effects, but regardless of the degree of caste‐specific extrinsic mortality. Mutations with antagonistic fitness effects within castes reduced lifespans of both castes, while mutations with between‐caste antagonistic effects decreased worker lifespans more than queen lifespans, and consequently increased lifespan divergences. Our results challenge the central explanatory role of extrinsic mortality for caste‐specific ageing in eusocial organisms and suggest that antagonistic pleiotropy affects castes differently due to reproductive monopolization by queens, hence, reproductive division of labor. Finally, these findings provide new insights into the evolution of tissue‐specific ageing in multicellular organisms in general.

COVID-19: the relationship between perceptions of risk and behaviours during lockdown

AimUnderstanding COVID-19 risk perceptions and their impact on behaviour can improve the effectiveness of public health strategies. Prior evidence suggests that, when people perceive uncontrollable risks to their health, they are less likely to engage in healthful behaviour. This article aims to understand the extent to which COVID-19 is perceived as an uncontrollable risk, and to assess whether this perceived risk is associated with health behaviour.Subject and methodsWe surveyed a nationally representative sample of 496 participants during the first UK lockdown. We assessed perceptions of COVID-19-related risk, self-reported adherence to infection control measures recommended by the UK Government, and general health behaviours. We predicted that increased perceived extrinsic mortality risk (the portion of mortality risk perceived to be uncontrollable) would disincentivise healthy behaviour.ResultsPerceived threat to life was the most consistent predictor of reported adherence to infection control measures. Perceived extrinsic mortality risk was found to have increased due to the pandemic, and was associated with lower reported adherence to Government advice on diet, physical activity, and smoking.ConclusionsOur findings suggest that health messages that highlight threat to life may be effective in increasing adherence to infection control, but may also lead to a reduction in health-promoting behaviours. We suggest that messages that highlight threat to life should be accompanied by statements of efficacy. Further, messages evoking feelings of concern for others may be effective in promoting compliance with anti-infection measures, without the potential for the unwelcome side-effect of discouraging healthy behaviour.

Intraspecific variation in incubation behaviours along a latitudinal gradient is driven by nest microclimate and selection on neonate quality

Abstract The strategies by which animals allocate reproductive effort across their lifetimes vary, and the causes of variation in those strategies are actively debated. In birds, most research has focused heavily on variation in clutch size and fecundity, but incubation behaviour and other functionally related traits have received less attention. Variation in incubation period duration is notable because time‐dependent sources of clutch mortality should impose strong directional selection to minimize the incubation period. However, life‐history theory predicts multiple mechanisms by which inter‐ and intraspecific variation in incubation behaviours may be adaptive. We conducted one of the first studies of intraspecific latitudinal variation in avian incubation behaviours across a large portion of a single species’ range. We placed motion‐activated nest cameras inside burrowing owl nests at five study sites to quantify variation in daily nest attentiveness, cumulative nest attendance and incubation period duration. We tested predictions of two alterative hypotheses that have been proposed to explain variation in incubation periods: the parental risk tolerance hypothesis and the neonate quality hypothesis. Daily nest attentiveness, cumulative nest attendance and incubation period duration in burrowing owls were all positively correlated with latitude. Burrowing owls reduced their daily nest attentiveness at low latitudes and on days when the average nest temperature was within the range that is optimal for embryo development. Further, longer incubation periods were most strongly associated with greater cumulative nest attendance instead of reduced daily nest attentiveness. These results support predictions of the neonate quality hypothesis: longer incubation periods result from stronger selection on neonate quality rather than selection to reduce reproductive effort in response to low extrinsic mortality risk. However, some owls facultatively reduced their daily nest attentiveness, and this result supports the general hypothesis that incubation decisions reflect a trade‐off between reproduction and self‐maintenance, and that the optimal solution to that trade‐off varies systematically in response to latitudinal gradients in adult mortality. A free Plain Language Summary can be found within the Supporting Information of this article.

Disease History and Life History Predict Behavioral Control of the COVID-19 Pandemic.

It is puzzling why countries do not all implement stringent behavioral control measures to prevent the spread of COVID-19 even though preventive behaviors have been proven to be the only effective means to stop the pandemic. We provide a novel evolutionary life history explanation whereby pathogenic and parasitic prevalence represents intrinsic rather than extrinsic mortality risk that drives slower life history strategies and the related disease control motivation in all animals but especially humans. Our theory was tested and supported based on publicly available data involving over 150 countries. Countries having a higher historical prevalence of infectious diseases are found to adopt slower life history strategies that are related to prompter COVID-19 containment actions by the government and greater compliance by the population. Findings could afford governments novel insight into the design of more effective COVID-19 strategies that are based on enhancing a sense of control, vigilance, and compliance in the general population.

Activity Patterns and Age-dependent Changes in Behavior in the Clonal Ant Platythyrea punctata

Age-based polyethism, wherein young individuals perform tasks within the nest and later transition to outside activities, is common among many social insects. This division of labor ensures the safety of workers with longer residual lifespans, such as brood caring nurses, and exposes only old individuals to increased extrinsic mortality risks. It appears however, that lifespan in workers is also shaped by intrinsic factors, such as senescence. The transition from nurse to forager is accompanied by various physiological and behavioral changes and seems to represent a crucial step in determining the subsequent life history of the individual. Here we investigate the daily activity pattern and detailed division of labor in colonies of the clonal ant Platythyrea punctata to better understand the dynamics of age polyethism in this peculiar species. We found that P. punctata colonies were mostly active during the day. At night, few workers were observed out of their nests. Workers showed a decrease in brood care and aggressive behavior over time. With increasing age individuals spent more time outside the nest and were more frequently observed walking. However, the transition to outside tasks did not follow a consistent time pattern in all colonies. Workers transitioning at a younger age seemed to die earlier than workers that began to forage later in life.

Integrating Mortality Risk and the Adaptiveness of Hibernation.

Low mortality rate is often associated with slow life history, and so far, has mainly been assessed through examinations of specific adaptations and lifestyles that limit mortality risk. However, the organization of activity time budgets also needs to be considered, since some activities and the time afforded for performing them may expose animals to higher mortality risks such as increased predation and/or increased metabolic stress. We examined the extent of activity time budgets contribution to explaining variation in life history traits in mammals. We specifically focused on hibernating species because of their marked seasonal cycle of activity/inactivity associated with very different mortality risks. Hibernation is considered a seasonal adaptation to prolonged periods of food shortage and cold. This inactivity period may also reduce both extrinsic and intrinsic mortality risks, by decreasing exposure to predators and drastically reducing metabolic rate. In turn, reduction in mortality may explain why hibernators have slower life history traits than non-hibernators of the same size. Using phylogenetically controlled models, we tested the hypothesis that longevity was positively correlated with the hibernation season duration (the time spent between immergence and emergence from the hibernaculum or den) across 82 different mammalian species. We found that longevity increased significantly with hibernation season duration, an effect that was particularly strong in small hibernators (<1.5 kg) especially for bats. These results confirm that hibernation not only allows mammals to survive periods of energy scarcity, but further suggest that activity time budgets may be selected to reduce mortality risks according to life history pace.

The behavioural constellation of deprivation: Causes and consequences

Socioeconomic differences in behaviour are pervasive and well documented, but their causes are not yet well understood. Here, we make the case that a cluster of behaviours is associated with lower socioeconomic status (SES), which we call "the behavioural constellation of deprivation." We propose that the relatively limited control associated with lower SES curtails the extent to which people can expect to realise deferred rewards, leading to more present-oriented behaviour in a range of domains. We illustrate this idea using the specific factor of extrinsic mortality risk, an important factor in evolutionary theoretical models. We emphasise the idea that the present-oriented behaviours of the constellation are a contextually appropriate response to structural and ecological factors rather than a pathology or a failure of willpower. We highlight some principles from evolutionary theoretical models that can deepen our understanding of how socioeconomic inequalities can become amplified and embedded. These principles are that (1) small initial disparities can lead to larger eventual inequalities, (2) feedback loops can embed early-life circumstances, (3) constraints can breed further constraints, and (4) feedback loops can operate over generations. We discuss some of the mechanisms by which SES may influence behaviour. We then review how the contextually appropriate response perspective that we have outlined fits with other findings about control and temporal discounting. Finally, we discuss the implications of this interpretation for research and policy.

Health behaviour, extrinsic risks, and the exceptions to the rule

Pepper & Nettle make a compelling case for how evolutionary thinking can help explain behaviours that cluster with deprivation. The role of extrinsic mortality risk in driving behaviour is probably important, but strong evidence is still lacking. By thinking carefully about behaviours seemingly at odds with an evolutionary life history perspective, we can gain important insights that will help refine theory.

Growth trajectories in wild geladas (Theropithecus gelada).

Life history and socioecological factors have been linked to species-specific patterns of growth across female vertebrates. For example, greater maternal investment in offspring has been associated with more discrete periods of growth and reproduction. However, in primates it has been difficult to test such hypotheses because very few studies have obtained growth measurements from wild populations. Here we utilize a promising noninvasive photogrammetric method-parallel lasers-to examine shoulder-rump (SR) growth in a wild primate, the gelada (Theropithecus gelada, Simien Mountains National Park, Ethiopia). In this species, a graminivorous diet coupled with high extrinsic infant mortality risk suggests that maternal investment in neonates is low. Therefore, in contrast with other closely related papionins, we expected female geladas to exhibit less discrete periods of growth and reproduction. For both sexes, we compared size-for-age patterns (N = 154 females; N = 110 males) and changes in growth velocity relative to major life history milestones. Female geladas finished 88.5% of SR growth by first sexual swelling, and 97.2% by first reproduction, reaching adult body size by 7.72 years of age. Compared to closely related papionins, gelada females finished more growth by first reproduction, despite producing relatively small, and presumably "cheap," neonates. Male geladas finished 85.4% of growth at dispersal, and 96.0% at estimated first birth. Contrary to other polygynous primates, males are larger than females because they grow for a longer period of time (not because they grow faster), surpassing females around 6 years of age when female growth slows. Our results demonstrate that parallel lasers are an easy and promising new method that can be used to construct comprehensive life history perspectives that were once out of reach for wild populations. Am. J. Primatol. 78:707-719, 2016. © 2016 Wiley Periodicals, Inc.

Trade-offs between acquired and innate immune defenses in humans

Immune defenses provide resistance against infectious disease that is critical to survival. But immune defenses are costly, and limited resources allocated to immunity are not available for other physiological or developmental processes. We propose a framework for explaining variation in patterns of investment in two important subsystems of anti-pathogen defense: innate (non-specific) and acquired (specific) immunity. The developmental costs of acquired immunity are high, but the costs of maintenance and activation are relatively low. Innate immunity imposes lower upfront developmental costs, but higher operating costs. Innate defenses are mobilized quickly and are effective against novel pathogens. Acquired responses are less effective against novel exposures, but more effective against secondary exposures due to immunological memory. Based on their distinct profiles of costs and effectiveness, we propose that the balance of investment in innate versus acquired immunity is variable, and that this balance is optimized in response to local ecological conditions early in development. Nutritional abundance, high pathogen exposure and low signals of extrinsic mortality risk during sensitive periods of immune development should all favor relatively higher levels of investment in acquired immunity. Undernutrition, low pathogen exposure, and high mortality risk should favor innate immune defenses. The hypothesis provides a framework for organizing prior empirical research on the impact of developmental environments on innate and acquired immunity, and suggests promising directions for future research in human ecological immunology.

Perceived extrinsic mortality risk and reported effort in looking after health: testing a behavioral ecological prediction.

Socioeconomic gradients in health behavior are pervasive and well documented. Yet, there is little consensus on their causes. Behavioral ecological theory predicts that, if people of lower socioeconomic position (SEP) perceive greater personal extrinsic mortality risk than those of higher SEP, they should disinvest in their future health. We surveyed North American adults for reported effort in looking after health, perceived extrinsic and intrinsic mortality risks, and measures of SEP. We examined the relationships between these variables and found that lower subjective SEP predicted lower reported health effort. Lower subjective SEP was also associated with higher perceived extrinsic mortality risk, which in turn predicted lower reported health effort. The effect of subjective SEP on reported health effort was completely mediated by perceived extrinsic mortality risk. Our findings indicate that perceived extrinsic mortality risk may be a key factor underlying SEP gradients in motivation to invest in future health.

Increased mortality exposure within the family rather than individual mortality experiences triggers faster life-history strategies in historic human populations.

Life History Theory predicts that extrinsic mortality risk is one of the most important factors shaping (human) life histories. Evidence from contemporary populations suggests that individuals confronted with high mortality environments show characteristic traits of fast life-history strategies: they marry and reproduce earlier, have shorter birth intervals and invest less in their offspring. However, little is known of the impact of mortality experiences on the speed of life histories in historical human populations with generally higher mortality risk, and on male life histories in particular. Furthermore, it remains unknown whether individual-level mortality experiences within the family have a greater effect on life-history decisions or family membership explains life-history variation.In a comparative approach using event history analyses, we study the impact of family versus individual-level effects of mortality exposure on two central life-history parameters, ages at first marriage and first birth, in three historical human populations (Germany, Finland, Canada). Mortality experience is measured as the confrontation with sibling deaths within the natal family up to an individual's age of 15.Results show that the speed of life histories is not adjusted according to individual-level mortality experiences but is due to family-level effects. The general finding of lower ages at marriage/reproduction after exposure to higher mortality in the family holds for both females and males. This study provides evidence for the importance of the family environment for reproductive timing while individual-level mortality experiences seem to play only a minor role in reproductive life history decisions in humans.

Origins and Evolution of GenusHomo

Recent fossil and archaeological finds have complicated our interpretation of the origin and early evolution of genus Homo. Using an integrated data set from the fossil record and contemporary human and nonhuman primate biology, we provide a fresh perspective on three important shifts in human evolutionary history: (1) the emergence of Homo, (2) the transition between non-erectus early Homo and Homo erectus, and (3) the appearance of regional variation in H. erectus. The shift from Australopithecus to Homo was marked by body and brain size increases, a dietary shift, and an increase in total daily energy expenditure. These shifts became more pronounced in H. erectus, but the transformation was not as radical as previously envisioned. Many aspects of the human life history package, including reduced dimorphism, likely occured later in evolution. The extant data suggest that the origin and evolution of Homo was characterized by a positive feedback loop that drove life history evolution. Critical to this process were probably cooperative breeding and changes in diet, body composition, and extrinsic mortality risk. Multisystem evaluations of the behavior, physiology, and anatomy of extant groups explicitly designed to be closely proxied in the fossil record provide explicit hypotheses to be tested on future fossil finds.