Allometry of human fertility and energy use

The flux of energy and materials constrains all organisms, and allometric relationships between rates of energy consumption and other biological rates are manifest at many levels of biological organization. Although human ecology is unusual in many respects, human populations also face energetic constraints. Here we present a model relating fertility rates to per capita energy consumption rates in contemporary human nations. Fertility declines as energy consumption increases with a scaling exponent of ?1/3 as predicted by allometric theory. The decline may be explained by parental trade‐offs between the number of children and the energetic investment in each child. We hypothesize that the ?1/3 exponent results from the scaling properties of the networked infrastructure that delivers energy to consumers. This allometric analysis of human fertility offers a framework for understanding the demographic transition to smaller family sizes, with implications for human population growth, resource use and sustainability.     

Determinants of human population growth

The 20th century has seen unprecedented growth of the human population on this planet. While at the beginning of the century the Earth had an estimated 1.6 billion inhabitants, this number grew to 6.1 billion by the end of the century, and further significant growth is a near certainty. This paper tries to summarize what factors lie behind this extraordinary expansion of the human population and what population growth we can expect for the future. It discusses the concept of demographic transition and the preconditions for a lasting secular fertility decline. Recent fertility declines in all parts of the world now make it likely that human population growth will come to an end over the course of this century, but in parts of the developing world significant population growth is still to be expected over the coming decades. The slowing of population growth through declining birth rates, together with still increasing life expectancy, will result in a strong ageing of population age structure. Finally, this paper presents a global level systematic analysis of the relationship between population density on the one hand, and growth and fertility rates on the other. This analysis indicates that in addition to the well-studied social and economic determinants, population density also presents a significant factor for the levels and trends of human birth rates.     

The Future of Tropical Forest Species1

Deforestation and habitat loss are widely expected to precipitate an extinction crisis among tropical forest species. Humans cause deforestation, and humans living in rural settings have the greatest impact on extant forest area in the tropics. Current human demographic trends, including slowing population growth and intense urbanization, give reason to hope that deforestation will slow, natural forest regeneration through secondary succession will accelerate, and the widely anticipated mass extinction of tropical forest species will be avoided. Here, we show that the proportion of potential forest cover remaining is closely correlated with human population density among countries, in both the tropics and the temperate zone. We use United Nations population projections and continent‐specific relationships between both total and rural population density and forest remaining today to project future tropical forest cover. Our projections suggest that deforestation rates will decrease as population growth slows, and that a much larger area will continue to be forested than previous studies suggest. Tropical forests retracted to smaller areas during repeated Pleistocene glacial events in Africa and more recently in selected areas that supported large prehistoric human populations. Despite many caveats, these projections and observations provide hope that many tropical forest species will be able to survive the current wave of deforestation and human population growth. A strategy to preserve tropical biodiversity might include policies to improve conditions in tropical urban settings to hasten urbanization and preemptive conservation efforts in countries with large areas of extant forest and large projected rates of future human population growth. We hope that this first attempt inspires others to produce better models of future tropical forest cover and associated policy recommendations.     

Global estimates and projections of world populations

The United Nations 2002 Revision of the world population projections has radically reviewed the results of the preceding revision. The results obtained show that it is reasonalbe revision. The results obtained show that it is reasonable to dispel the fear of an excessive and perhaps uncontrolled growth of world population, which alarmed public opinion and many international organizations in the last decades. Even in the medium variant, by the year 2050 world population will be less than 9 billion, with zero growth. In view of the fact that fertility will remain below replacement level, after that a process of slow decrease might set in, despite a further sharp increase in survival rates. The objectives of the projections are of an administrative nature: firstly, to adapt global and local policies to the real needs of the population; secondly, to distinguish the trends that are favourable or harmful for the population's greater wel-being; thirdly, to govern society in such a way as to achieve the highest possible standard of living, which must be extended to everyone, long-lasting, and therefore compatible with the limits imposed by the environment; the final objective is not to subtract resources from the future generations.     

Population and prehistory I: Food-dependent population growth in constant environments

We present a demographic model that describes the feedbacks between food supply, human mortality and fertility rates, and labor availability in expanding populations, where arable land area is not limiting. This model provides a quantitative framework to describe how environment, technology, and culture interact to influence the fates of preindustrial agricultural populations. We present equilibrium conditions and derive approximations for the equilibrium population growth rate, food availability, and other food-dependent measures of population well-being. We examine how the approximations respond to environmental changes and to human choices, and find that the impact of environmental quality depends upon whether it manifests through agricultural yield or maximum (food-independent) survival rates. Human choices can complement or offset environmental effects: greater labor investments increase both population growth and well-being, and therefore can counteract lower agricultural yield, while fertility control decreases the growth rate but can increase or decrease well-being. Finally we establish equilibrium stability criteria, and argue that the potential for loss of local stability at low population growth rates could have important consequences for populations that suffer significant environmental or demographic shocks.     

Potential for Worldwide Displacement of Fossil-Fuel Electricity by Nuclear Energy in Three Decades Based on Extrapolation of Regional Deployment Data

There is an ongoing debate about the deployment rates and composition of alternative energy plans that could feasibly displace fossil fuels globally by mid-century, as required to avoid the more extreme impacts of climate change. Here we demonstrate the potential for a large-scale expansion of global nuclear power to replace fossil-fuel electricity production, based on empirical data from the Swedish and French light water reactor programs of the 1960s to 1990s. Analysis of these historical deployments show that if the world built nuclear power at no more than the per capita rate of these exemplar nations during their national expansion, then coal- and gas-fired electricity could be replaced worldwide in less than a decade. Under more conservative projections that take into account probable constraints and uncertainties such as differing relative economic output across regions, current and past unit construction time and costs, future electricity demand growth forecasts and the retiring of existing aging nuclear plants, our modelling estimates that the global share of fossil-fuel-derived electricity could be replaced within 25–34 years. This would allow the world to meet the most stringent greenhouse-gas mitigation targets.     

A population biological approach to the collective dynamics of countries undergoing demographic transition

Birth rates have been declining in higher-income countries since the middle of the 19th century. A growing number of other countries have entered this demographic transition to lower fertility, as socioeconomic development continues. Analyses of this demographic transition vary widely, but most analyze individual populations in isolation from others, and most come from fields outside the biological sciences. Here, we develop a population biological model of population dynamics in higher-income countries. Individual countries evolve through density-regulated growth, where gradual evolution toward higher population densities boosts productivity (and hence socioeconomic growth) through economics of agglomeration and scale, in turn reducing birth rates. The exchange of technology and capital between countries can further boost productivity gains in any given country, thus contributing to its demographic transition. As a result, countries can down-regulate one another's population growth through mutual improvements in productivity. The model is fitted to time series data on population size, GDP per capita, and birth rates for the United States, the United Kingdom, and France. The metapopulation dynamics are also characterized across a range of parameter values close to the fitted values. This work may help advance population biological approaches to understanding the implications of the fertility demographic transition for modern human populations. This is relevant to developing long-term predictions of the earth's total population size, which must be based upon a model that incorporates underlying mechanisms.     

Uncertainties in predicting the demographic impact of AIDS

A model giving the demographic impact of AIDS is analysed to examine the sensitivity of the projections when various complicating features are included. The model deals with age and sexual partner change rate as continuous variables and uses a device to specify arbitrary correlations between the ages of the people who form sexual partnerships. The device ensures consistency, in that the amount of partner formation is the same regardless of whether the partnerships are counted from the point of view of males or females. Arbitrary correlation between partner change rate and fertility is also permitted. The results show the uncertainty in model predictions that population growth will reduce over the next 20 years to approximately zero in parts of East Africa severely affected by the AIDS epidemic. The main sources of uncertainty in the model predictions are assumptions concerning the correlation between ages in a partnership, the correlation between partner change rate and fertility, the incubation period of AIDS, and the variability of the female partner change rate.     

Dimensions of global population projections: what do we know about future population trends and structures?

The total size of the world population is likely to increase from its current 7 billion to 8–10 billion by 2050. This uncertainty is because of unknown future fertility and mortality trends in different parts of the world. But the young age structure of the population and the fact that in much of Africa and Western Asia, fertility is still very high makes an increase by at least one more billion almost certain. Virtually, all the increase will happen in the developing world. For the second half of the century, population stabilization and the onset of a decline are likely. In addition to the future size of the population, its distribution by age, sex, level of educational attainment and place of residence are of specific importance for studying future food security. The paper provides a detailed discussion of different relevant dimensions in population projections and an evaluation of the methods and assumptions used in current global population projections and in particular those produced by the United Nations and by IIASA.     

A DEMOGRAPHIC TRANSITION ALTERED THE STRENGTH OF SELECTION FOR FITNESS AND AGE‐SPECIFIC SURVIVAL AND FERTILITY IN A 19TH CENTURY AMERICAN POPULATION

Modernization has increased longevity and decreased fertility in many human populations, but it is not well understood how or to what extent these demographic transitions have altered patterns of natural selection. I integrate individual‐based multivariate phenotypic selection approaches with evolutionary demographic methods to demonstrate how a demographic transition in 19th century female populations of Utah altered relationships between fitness and age‐specific survival and fertility. Coincident with this demographic transition, natural selection for fitness, as measured by the opportunity for selection, increased by 13% to 20% over 65 years. Proportional contributions of age‐specific survival to total selection (the complement to age‐specific fertility) diminished from approximately one third to one seventh following a marked increase in infant survival. Despite dramatic reductions in age‐specific fertility variance at all ages, the absolute magnitude of selection for fitness explained by age‐specific fertility increased by approximately 45%. I show that increases in the adaptive potential of fertility traits followed directly from decreased population growth rates. These results suggest that this demographic transition has increased the adaptive potential of the Utah population, intensified selection for reproductive traits, and de‐emphasized selection for survival‐related traits.     

Energetic demand of multiple dependents and the evolution of slow human growth

This study investigates the consequences of the human foraging niche and multiple dependent offspring on the optimal growth trajectory of humans. We test the hypothesis that the human pattern of slow human growth between age at weaning and puberty helps defer the compound energetic demand on parents with multiple dependents, by using growth and demographic data from two foraging societies, the Ache of eastern Paraguay and the Dobe Ju/'hoansi of Botswana and Namibia. We run simulations of observed and potential growth trajectories among sub-adults and their consequent energetic demands on parents given profiles of fertility, mortality, consumption and production. We find that either sub-adult production or food subsidies from other people must substantially increase in order to compensate for the dramatic increase in energetic demand on parents if offspring were to grow faster at younger ages. Our conclusion is that slow human growth followed by a rapid adolescent growth spurt may have facilitated rising human fertility rates and greater investments in neural capital.     

The heritability of fertility makes world population stabilization unlikely in the foreseeable future

The forecasting of the future growth of world population is of critical importance to anticipate and address a wide range of global challenges. The United Nations produces forecasts of fertility and world population every two years. As part of these forecasts, they model fertility levels in post-demographic transition countries as tending toward a long-term mean, leading to forecasts of flat or declining population in these countries. We substitute this assumption of constant long-term fertility with a dynamic model, theoretically founded in evolutionary biology, with heritable fertility. Rather than stabilizing around a long-term level for post-demographic transition countries, fertility tends to increase as children from larger families represent a larger share of the population and partly share their parents' trait of having more offspring. Our results suggest that world population will grow larger in the future than currently anticipated.     

The opening of Pandora’s Box: climate change impacts on soil fertility and crop nutrition in developing countries

Feeding the world’s growing population is a serious challenge. Food insecurity is concentrated in developing nations, where drought and low soil fertility are primary constraints to food production. Many crops in developing countries are supported by weathered soils in which nutrient deficiencies and ion toxicities are common. Many systems have declining soil fertility due to inadequate use of fertility inputs, ongoing soil degradation, and increasingly intense resource use by burgeoning populations. Climate models predict that warmer temperatures and increases in the frequency and duration of drought during the 21st century will have net negative effects on agricultural productivity. The potential effects of climate change on soil fertility and the ability of crops to acquire and utilize soil nutrients is poorly understood, but is essential for understanding the future of global agriculture. This paper explores how rising temperature, drought and more intense precipitation events projected in climate change scenarios for the 21st century might affect soil fertility and the mineral nutrition of crops in developing countries. The effects of climate change on erosion rates, soil organic carbon losses, soil moisture, root growth and function, root-microbe associations and plant phenology as they relate to mineral nutrition are discussed. Our analysis suggests that the negative impacts of climate change on soil fertility and mineral nutrition of crops will far exceed beneficial effects, which would intensify food insecurity, particularly in developing countries.     

Where next?

This paper provides a personal perspective on the rich discussions at the Bixby Forum. The size, rate of growth and age structure of the human population interact with many other key factors, from environmental change to governance. While the details of future interactions are sometimes difficult to predict, taken together they pose sombre threats to a socially and economically sustainable future for the rich and to any realistic possibility of lifting the world''s bottom two billion people out of poverty. Adaptive changes will be needed to cope with an ageing population in countries with low fertility or below, but these are achievable. More worrying, continued rapid population growth in many of the least developed countries could lead to hunger, a failure of education to keep pace with growing numbers, and conflict. The assumption that the demographic transition from high to low birth rates occurs as a result of exogenous social and economic forces is being replaced by a clearer understanding of the many barriers that separate women from the knowledge and technologies they need to manage their childbearing within a human rights framework. The forum ended with a clear consensus that much more emphasis needs to be given to meeting the need for family planning and to investing in education.     

Modeling neolithic dispersal in central Europe: demographic implications

On the basis of new examination of ancient DNA and craniometric analyses, Neolithic dispersal in Central Europe has been recently explained as reflecting colonization or at least a major influx of near eastern farmers. Given the fact that Neolithic dispersal in Central Europe was very rapid and extended into a large area, colonization would have to be associated with high population growth and fertility rates of an expanding Neolithic population. We built three demographic models to test whether the growth and fertility rates of Neolithic farmers were high enough to allow them to colonize Central Europe without admixture with foragers. The principle of the models is based on stochastic population projections. Our results demonstrate that colonization is an unlikely explanation for the Neolithic dispersal in Central Europe, as the majority of fertility and growth rate estimates obtained in all three models are higher than levels expected in the early Neolithic population. On the basis of our models, we derived that colonization would be possible only if (1) more than 37% of women survived to mean age at childbearing, (2) Neolithic expansion in Central Europe lasted more than 150 years, and (3) the population of farmers grew in the entire settled area. These settings, however, represent very favorable demographic conditions that seem unlikely given current archaeological and demographic evidence. Therefore, our results support the view that Neolithic dispersal in Central Europe involved admixture of expanding farmers with local foragers. We estimate that the admixture contribution from foragers may have been between 55% and 72%.     

Bevölkerungswachstum: Bildung ist die Lösung

Population growth – education is the answer Over the last decades world population has been constantly growing by some 80 million per year. Whereas the growth rate as well as the fertility rate have been cut by half since the 1970th, population growth will continue well over mid‐century. As the developed countries have completed the demographic transition from high mortality and fertility rates to low ones, population growth is fading out there or has already been reversed into decline. In the least developed countries mortality has fallen as well, whereas fertility decline has stalled. Therefor population growth is very high making the solution of the widespread problems in this part of the world more and more difficult. One obvious way out of this trap would be a better education that could open new development perspectives. A positive side effect is that educated women have much less offspring than their counterparts who never went to school.     

Bioethanol and biodiesel: Alternative liquid fuels for future generations

The global population is expected to increase by approximately 3 billion people by 2050. With this increase in population, industry, transportation the cost of fossil fuels will grow dramatically. New technologies are needed for fuel extraction using feedstocks that do not threaten food security, cause minimal or no loss of natural habitat and soil carbon. At the same time, waste management has to be improved and environmental pollution should be minimized or eliminated. Liquid biofuels such as lignocellulosic‐based ethanol from plant biomass and algal‐based biodiesel are sustainable, alternative biofuels that could stabilize national security and provide clean energy for future generations. Ideally, the technology should also foster recycling of agricultural feedstocks and improve soil fertility and human health. This article provides updated information on the energy potential and breadth of liquid biofuel biotechnology.     

Effects of the demographic transition on the genetic variances and covariances of human life‐history traits

The recent demographic transitions to lower mortality and fertility rates in most human societies have led to changes and even quick reversals in phenotypic selection pressures. This can only result in evolutionary change if the affected traits are heritable, but changes in environmental conditions may also lead to subsequent changes in the genetic variance and covariance (the G matrix) of traits. It currently remains unclear if there have been concomitant changes in the G matrix of life‐history traits following the demographic transition. Using 300 years of genealogical data from Finland, we found that four key life‐history traits were heritable both before and after the demographic transition. The estimated heritabilities allow a quantifiable genetic response to selection during both time periods, thus facilitating continued evolutionary change. Further, the G matrices remained largely stable but revealed a trend for an increased additive genetic variance and thus evolutionary potential of the population after the transition. Our results demonstrate the validity of predictions of evolutionary change in human populations even after the recent dramatic environmental change, and facilitate predictions of how our biology interacts with changing environments, with implications for global public health and demography.     

Comparison of linear and nonlinear models for human population dynamics

In standard demographic practice, population projections are commonly based on one-sex linear models of the Lotka-Leslie type. We demonstrate here that such projections based solely on time-invariant, age-specific male fertilities and mortalities are incompatible with those based solely on female fertilities and mortalities. This incompatibility obtains even in the singular case where effective male and female fertility functions are equal, and generate equal ultimate rates of growth. In standard demographic practice, the incompatibility is initially masked, since one-sex fertility functions are generally calculated from the same initial-time data and thereby tautologically forced to initially concur; however, with the passage of any finite time, the incompatibility reasserts itself—the only exception being the uninteresting case where the system is already in the stationary age and sex distribution of balanced exponential growth.An example is adduced of a nonlinear (age-free) system whose true rate of ultimate growth is correctly bracketed by the male and female one-sex rates of ultimate growth. Analysis of more general two-sex models shows that the two one-sex growth rates, calculated for arbitrary male and female initial age distributions, need not bracket the true rate. We show, however, that models exist such that with appropriate choices of initial conditions, this bracketing will occur.     

A life‐history perspective on the demographic drivers of structured population dynamics in changing environments

Current understanding of life‐history evolution and how demographic parameters contribute to population dynamics across species is largely based on assumptions of either constant environments or stationary environmental variation. Meanwhile, species are faced with non‐stationary environmental conditions (changing mean, variance, or both) created by climate and landscape change. To close the gap between contemporary reality and demographic theory, we develop a set of transient life table response experiments (LTREs) for decomposing realised population growth rates into contributions from specific vital rates and components of population structure. Using transient LTREs in a theoretical framework, we reveal that established concepts in population biology will require revision because of reliance on approaches that do not address the influence of unstable population structure on population growth and mean fitness. Going forward, transient LTREs will enhance understanding of demography and improve the explanatory power of models used to understand ecological and evolutionary dynamics.