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This chapter describes a formal demographic perspective on human population dynamics. It first attempts to summarise the way in which human population dynamics are treated in the more technical and theoretical demographic literature. The next section considers some demographic fundamentals, including population structure (especially the age and sex composition) and the three components of population change: fertility, mortality and migration. The third part looks at some of the formal models which demographers have developed to help understand population change. These models make several assumptions in order to simplify a complex reality. One of these is that migration is zero: populations with zero migration are said to be closed. An attraction of this is that, if migration can be ignored, simple relationships exist between fertility, mortality, the population growth rate and the age structure.
In the fourth section population dynamics in the short to medium term are considered. The age and sex structure of a population is itself a dynamic feature, containing a record of the population's past fertility, mortality and migration. Moreover, the future age–sex structure is determined by past and current events. Discussion of these aspects of population dynamics leads naturally in the fifth section to a consideration of population momentum, or what is sometimes, inaccurately, called the ‘demographic time bomb’. The origins of population momentum are explained.
Finally, in the sixth and seventh sections long-run population dynamics are explored in the context of the demographic transitions in Europe and contemporary Africa.
By
Helen MacBeth, Honorary Research Fellow Department of Anthropology, School of Social Sciences and Law, Oxford Brookes University, Headington, Oxford OX3 0BP, U.K.,
Paul Collinson, Honorary Research Fellow Department of Anthropology, School of Social Sciences and Law, Oxford Brookes University, Headington, Oxford OX3 0BP, U.K.
Evolutionary biology became gradually transformed through the twentieth century from what was termed ‘typological thinking’ to ‘population thinking’. This happened from the increasing recognition that differences between individuals were as important as similarities. Variation contains the raw material of evolution and variation can exist only in populations. The transformation harmonised the relationship of evolutionary biology with other population based biologies: genetics, ecology and epidemiology, for example. More significantly, however, in humans it also established potential connections with some of the social sciences and especially sociology, social anthropology, human geography and social psychology, all of which are fundamentally population based areas of knowledge.
Population can thus be seen as a bridge between the ‘two cultures’ of natural sciences and the humanities. This was most effectively recognised by J.W.S. Pringle at Oxford, who was instrumental in establishing there the field of Human Sciences, essentially based on the analysis of human population structures from all perspectives. In the past twenty-five years the field has developed dramatically.
The concept of population is not, however, one without difficulties. Even the definition of a population can be a major problem. Human groups rarely exist as discrete, more or less uniform entities even at any one time, and they have never existed over time. Typically they intergrade, often very gradually. Then while one most frequently thinks of populations in some spatial context, they also exist in vertical dimensions such as social class and caste, and in ecological and economic terms reflecting natural environmental heterogeneity.
The complex processes of human population dynamics work themselves out, not in an abstract limbo but, as for other species, in particular environmental settings. The specific physical and biotic characteristics of such settings may crucially affect these dynamics and vice versa. From their local environments, and/or from any others to which personal travel and exchange systems allow access, people must acquire the material resources, such as food and water, necessary for individual and population survival. Complicating factors peculiar to Homo sapiens have not abolished this fundamental dependence on our environments; but they have both changed the nature and time-scale of that dependence, and made its reality harder to see.
One of these complications, with a major effect on human population–environment relations, is our capacity to reach beyond the local environment to obtain resources. In a now globalised economy, access to other environments' resources is scarcely impeded by geographical barriers or distance, though very much impeded by poverty. Even in earlier times and more straitened circumstances, flows of material necessities have commonly extended well beyond the settings local to particular communities. The potential implications for population dynamics are huge.
Other complications apply in human societies too, even within local settings. People view landscapes and resources through culturally informed spectacles. They imbue them with symbolic meanings, religious beliefs, emotional attachments, proprietorial claims, and senses of individual and collective identity. They develop and communicate sophisticated understandings of geography, seasonality and human impacts.
By
Helen MacBeth, Honorary Research Fellow Department of Anthropology, School of Social Sciences and Law, Oxford Brookes University, Headington, Oxford OX3 0BP, U.K.,
Paul Collinson, Honorary Research Fellow Department of Anthropology, School of Social Sciences and Law, Oxford Brookes University, Headington, Oxford OX3 0BP, U.K.
Humans can be studied from a variety of academic perspectives. In some biological disciplines the focus may be on parts of individuals, perhaps even molecular parts, while in others humans are considered zoologically as an entire species, Homo sapiens, in the order Primatii. Some biologists are concerned with the physiologically functioning body of an individual, while others are interested in world-wide human diversity, distribution and population groupings. Among the social sciences, the focus is also frequently on the group, but here the emphasis tends to be upon the social and cultural factors which underpin the way such phenomena as societies, communities and ethnic groups are constructed, delimited and defined. There are also the behavioural sciences, which utilise both biological and social information in relation to the study of the group, the individual or elements in the behaviour of the individual. Recognition of the number of perspectives on these population variables emphasises the need to study material across the boundaries of traditional academic disciplines and several of these perspectives are introduced in this volume.
The twentieth century began with observers of the human condition integrating, in what we now recognise to be a confused way, their ideas on the biological and social nature of humans. During the century the fragmentation of all disciplines grew, something exemplified most particularly in the divergence between biological and social pursuits of knowledge.
The beginning of the twenty-first century is a most opportune time to look back fifty years to see what has happened to the spatial patterns of world population distribution, and to try to look forward over a similar period to see what is likely to happen, especially as we are in the midst of an almost inexorable process of massive concentration of world population. That concentration must be examined within the context of other enormous demographic changes that have taken place since 1950 and are likely to take place before 2050. Naturally, the reliability of the retrospective analysis is greater than that of the prospective analysis, although I am relieved to note that most of my speculations and predictions made in comparable papers about three decades ago (Clarke 1971, 1972) have not been far from the mark. As Hinde (Chapter 2) has mentioned, the perspectives of demographers and geographers overlap, but the latter pay great attention to spatial distribution and migration.
Rapid world population growth
The mid-twentieth century was a demographic threshold when world population growth accelerated. At that time, the world population was just over 2.5 billion; according to the United Nations Population Division it reached 6 billion in October 1999. It had taken roughly a century and a half for world population to multiply two and a half times from 1 billion to 2.5 billion, but in the second half of the twentieth century the world population grew so rapidly that 3.5 billion were added to the world total, one billion being added during the last 12 years (Table 3.1).
By
L.B. Jorde, Department of Human Genetics, University of Utah Health Sciences Center, Salt Lake City, UT 84112, U.S.A.,
M.J. Bamshad, Department of Pediatrics, University of Utah Health Sciences Center, Salt Lake City, UT 84112, U.S.A.,
W.S. Watkins, Department of Human Genetics, University of Utah Health Sciences Center, Salt Lake City, UT 84112, U.S.A.,
C.E. Ricker, Department of Human Genetics, University of Utah Health Sciences Center, Salt Lake City, UT 84112, U.S.A.,
M.E. Dixon, Department of Human Genetics, University of Utah Health Sciences Center, Salt Lake City, UT 84112, U.S.A.,
B.B. Rao, Department of Anthropology, Andhra University, Visakhapatnam, Andhra Pradesh, India,
B.V.R. Prasad, Department of Anthropology, Andhra University, Visakhapatnam, Andhra Pradesh, India,
J.M. Naidu, Department of Anthropology, Andhra University, Visakhapatnam, Andhra Pradesh, India
The Indian subcontinent, which contains nearly one-sixth of the world's population, has received migrants from many sources during its history. Its population is consequently diverse, both culturally and genetically (Nei and Roychoudhury 1982; Majumder and Mukherjee 1993; Majumder 1998). The patterns of this diversity can offer important clues to evolutionary history. In addition, unique sociocultural phenomena in India have influenced mating patterns for thousands of years. It is expected that these patterns have in turn affected the genetic structure of the population. For these reasons, the Indian population is an appealing subject for population genetic analysis. This chapter, while concentrating on some populations in southern India, shows how modern genetic techniques can play an important part in our understanding of human population dynamics.
Three major waves of immigration have influenced the genetic structure of India. Palaeolithic immigrants are thought to have originated from Africa, or possibly Australia, and may have contributed to the gene pool of present-day tribal populations (Maloney 1974; Chandler 1988; Majumder and Mukherjee 1993; Cavalli-Sforza et al. 1994). A second major wave of proto-Dravidian-speaking immigrants came from the Fertile Crescent area c. 10,000 years ago and populated most of the subcontinent (Cavalli-Sforza et al. 1994). The third major wave, emanating from west-central Asia about 3,500 years ago, consisted of Indo-European-speaking ‘Caucasoid’ populations. This immigration event is marked archaeologically by the appearance of Painted Grey Ware (Thapar 1980).
This chapter compares causes and consequences of population growth in the Lua' and the Hmong highland minority groups in Southeast Asia. The study illustrates roles of cultural patterns in, and impacts of, modernisation on population dynamics. It exemplifies many of the processes introduced in earlier chapters in relation to demography, change in social and health conditions, and population genetics. It relates changes in mortality, fertility and migration, and in geographic and social boundaries, to socioeconomic modernisation, including public health programmes, education, development and the market economy. This condensation of more detailed papers by Kunstadter and colleagues (see References section) suggests the value of holistic and longitudinal approaches.
Data were collected over four decades by conventional anthropological participant observation, health examinations, unstructured and structured interviews, surveys and questionnaires, some modelled on those developed for international comparative studies, such as the Demographic and Health Survey, with appropriate modifications based on discussions with members of the populations. Native speakers conducted most surveys.
Determinants of primary population variables
In Lua' and Hmong populations almost all fertility is confined to marriage. Thus age at marriage, proportion who marry, and divorce rates are important determinants of fertility, as are use of deliberate methods to delay, space or end pregnancies (Davis and Blake 1956).
Important biological determinants of fertility include age of reproductive maturity and senescence. These are influenced by nutritional status and infections, especially sexually transmitted diseases. Breastfeeding suppresses ovulation and protects infants from infectious diseases.
By
Jaume Bertranpetit, Unitat de Biologia Evolutiva, Facultat de Ciencies de la Salut i de la Vida, Universitat Pompeu Fabra, Doctor Aiguader 80, 08003 Barcelona, Spain,
Francesc Calafell, Unitat de Biologia Evolutiva, Facultat de Ciencies de la Salut i de la Vida, Universitat Pompeu Fabra, Doctor Aiguader 80, 08003 Barcelona, Spain
Humans are biological organisms. After chapters on demographic and socioeconomic perspectives on human populations and their spatial distribution, it is now appropriate to focus on contemporary ways in which world-wide human biological diversity and dynamics can be studied through human population genetics in its development in the era of the Human Genome Project.
The description of human differences is ancient. There are references in antiquity to humans with different physical features, such as those described by Herodotus, but the confusion between biological features and those we would today consider cultural was common until the twentieth century and still persists in some material. Over the course of the last century, there have been continuous debates about human diversity, but these debates were dominated by descriptions of morphological characteristics, for example pigmentation, and body shape and size. These descriptions in turn gave rise to ‘racial’ classifications, which remain highly significant socially. Even with the increase in understanding of genetic processes, much of this body of work has done nothing to further the understanding of human diversity, because in most cases the analyses did not take into consideration two basic issues.
The first issue is the evolutionary perspective on a given pattern of diversity. Following a tradition in the description of nature itself, the description of human populations continued to take, even recently, a naturalistic perspective, more in accordance with a pre-Darwinian view of detailed description than with its meaning in biological terms.
By
Helen MacBeth, Honorary Research Fellow Department of Anthropology, School of Social Sciences and Law, Oxford Brookes University, Headington, Oxford OX3 0BP, U.K.,
Paul Collinson, Honorary Research Fellow Department of Anthropology, School of Social Sciences and Law, Oxford Brookes University, Headington, Oxford OX3 0BP, U.K.
It is clear that there are many perspectives on human populations and that these are studied within different disciplines, whose terminology and literature are becoming increasingly specialised. Furthermore, the features and conditions of the populations are always changing and the processes of this are viewed and reported upon from different angles. What is required, therefore, for a complete understanding of the complex dynamics of human populations is an appreciation of the work of specialists in different disciplines, something which this volume provides. It is anticipated that the book will be useful to all who consider human populations in their research or their studies, and particularly those who are undertaking degree courses in human sciences, anthropology, demography, human geography, ecology and human biology. The book will also have a wider appeal for the more general reader who wishes to advance their understanding of human population dynamics by adopting a wide-ranging disciplinary focus.
This volume arises from the fifteenth workshop of the Biosocial Society, which was held at the Pauling Human Sciences Centre, Oxford, in 2000. The Society thanks Ros Odling-Smee and the Centre for accommodating the conference. The help with arrangements for and at the workshop by the team of postgraduates from Oxford Brookes {University} is gratefully acknowledged, and the Society is thanked for providing finances for the workshop. The Editors also wish to thank Chris McDonaugh for reading the book and making helpful suggestions, Jennifer Jay for work on the glossary, index and referencing, and finally Geoff Harrison for writing the Foreword.
By
Emily K. Rousham, Department of Human Sciences, Loughborough University, Loughborough, Leics LE11 3TU, U.K.,
Louise T. Humphrey, Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW17 5BD, U.K.
Changes in child survival rates are very significant in the dynamics of human populations, and mortality is an important variable in studies of demography, genetics and anthropology. Furthermore, rates of child survival provide an important indication of the general biological welfare of a population.
Children are a particularly vulnerable group, both physically and socially, within human populations (Caldwell 1996). This vulnerability is reflected in mortality rates, which are highest during infancy, decrease from 1 to 4 years and reach a minimum from 5 to 9 years of age. The physical vulnerability of children stems from an immature immune system, rendering them particularly susceptible to infectious diseases. This is coupled with a high energy and nutrient requirement relative to body weight. These intakes are required to fuel the high energy demands of the human brain, as well as for somatic growth and development. One of the most critical periods for child survival is the weaning period, typically from six months onwards, when infants make the transition from dependence on mother's milk to reliance on the local diet (Martorell 1995). Children are also socially vulnerable, being entirely dependent on adult carers for the first year of life and remaining highly dependent up to the age of five. This social dependence means that children are susceptible to conscious or unconscious neglect (Caldwell 1996).
This chapter examines the sociocultural and biological correlates of child survival, contrasting populations with low and high survival rates.
Most aphids are cyclic parthenogens, so are ideal models in studies of the mechanisms and consequences of sex and recombination. However, owing to a shortage of physical and genetic markers, there have been few studies of the most fundamental genetic processes in these organisms. For example, it is not known whether autosomal segregation during male spermatogenesis is in Mendelian proportions: we address that question here. The aphid Myzus persicae has a typical karyotype of 2n = 12 in females (XX), while males are XO (2n = 11). During male meiosis, only the spermatocytes with an X chromosome are viable. We hypothesized that assortment of autosomes might be non-random because chromosomal imprinting leading to elimination of the paternal autosomes is seen in the closely related coccoids. In other aphid models, we have observed segregation distortions at single microsatellite loci (Wilson, 2000). Such distortions may have nothing to do with ‘selfish’ behaviour, but may be caused by mutation accumulation causing fitness differentials. Thus single-locus distortions might be predicted to be more likely to be detected via the male lines of clones that have lost the ability to reproduce sexually (male-producing obligate parthenogenesis (androcyclic)). Using microsatellites we show that genetic imprinting or selfish autosome behaviour does not occur in male M. persicae. Generally, loci segregated in Mendelian proportions in both sexes of cyclically parthenogenetic (holocyclic) clones. However, in androcyclic clones, segregation distortions consistently involved the same two autosomes. This is consistent with linkage of markers to deleterious mutations associated with a loss of sexual reproduction.
We investigate how sampling of parents or children based on their extreme phenotypic values selected from clinical databases would affect the power of identification of quantitative trait loci (QTL) by a transmission disequilibrium test (TDT). We consider three selective sampling schemes based on the selection of phenotypic values of parents or children in nuclear families: (1) two children, one of extreme value, the other random; (2) two children extremely discordant; (3) one parent of extreme value. Other family members not specified will be recruited randomly with regard to phenotypic values. Our study shows that the second sampling scheme can always enhance the power for QTL identification, sometimes dramatically so. The increase in the statistical power of the TDT is particularly dramatic when h2 at the QTL under test is small or intermediate (e.g. 0·05 or 0·10). For the other two sampling schemes, under dominant effects at the QTL, the power is always increased relative to random sampling; however, under recessive or additive genetic effects, the power gain is generally minor or even decreased a little sometimes. Allele frequencies at the QTL and the selection stringency are important for determining the effect of selective sampling on the power of QTL identification. Our study is useful as a practical guideline on how to perform the TDT efficiently in practice by taking advantage of the extensive databases accumulated that are enriched with people of extreme phenotypic values.
The co-occurrence of three chromosome-wide phenomena – imprinting, facultative heterochromatization and diffuse centromere – in the mealybug Planococcus lilacinus makes investigation of the genomics of this species an attractive prospect. In order to estimate the complexity of the genome of this species, 300 random stretches of its DNA, constituting ∼0·1% of the genome, were sequenced. Coding sequences appear to constitute ∼53·5%, repeat sequences ∼44·5% and non-coding single-copy sequences ∼2% of the genome. The proportion of repetitive sequences in the mealybug is higher than that in the fruit fly Drosophila melanogaster (∼30%). The mealybug genome (∼220 Mb) is about 1·3 times the size of the fly genome (∼165 Mb) and its GC content (∼35%) less than that of the fly genome (∼40%). The relative abundance of various dinucleotides, as analysed by the method of Gentles and Karlin, shows that the dinucleotide signatures of the two species are moderately similar and that in the mealybug there is neither over-representation nor under-representation of any dinucleotide.
Natural populations are structured spatially into local populations and genetically into diverse ‘genetic backgrounds’ defined by different combinations of selected alleles. If selection maintains genetic backgrounds at constant frequency then neutral diversity is enhanced. By contrast, if background frequencies fluctuate then diversity is reduced. Provided that the population size of each background is large enough, these effects can be described by the structured coalescent process. Almost all the extant results based on the coalescent deal with a single selected locus. Yet we know that very large numbers of genes are under selection and that any substantial effects are likely to be due to the cumulative effects of many loci. Here, we set up a general framework for the extension of the coalescent to multilocus scenarios and we use it to study the simplest model, where strong balancing selection acting on a set of n loci maintains 2n backgrounds at constant frequencies and at linkage equilibrium. Analytical results show that the expected linked neutral diversity increases exponentially with the number of selected loci and can become extremely large. However, simulation results reveal that the structured coalescent approach breaks down when the number of backgrounds approaches the population size, because of stochastic fluctuations in background frequencies. A new method is needed to extend the structured coalescent to cases with large numbers of backgrounds.
Epistatic variance can be an important source of variation for complex traits. However, detecting epistatic effects is difficult primarily due to insufficient sample sizes and lack of robust statistical methods. In this paper, we develop a Bayesian method to map multiple quantitative trait loci (QTLs) with epistatic effects. The method can map QTLs in complicated mating designs derived from the cross of two inbred lines. In addition to mapping QTLs for quantitative traits, the proposed method can even map genes underlying binary traits such as disease susceptibility using the threshold model. The parameters of interest are various QTL effects, including additive, dominance and epistatic effects of QTLs, the locations of identified QTLs and even the number of QTLs. When the number of QTLs is treated as an unknown parameter, the dimension of the model becomes a variable. This requires the reversible jump Markov chain Monte Carlo algorithm. The utility of the proposed method is demonstrated through analysis of simulation data.
Clines for size and stress resistance traits have been described for several Drosophila species and replicable clines across different species may indicate climatic selection. Here we consider clines in stress resistance traits in an Australian endemic species, D. serrata, by comparing levels of variation within and among isofemale lines initiated with flies collected from the eastern coast of Australia. We also consider clinal variation in chill coma recovery, a trait that has recently been shown to exhibit high levels of variation among Drosophila species. Patterns were compared with those in the cosmopolitan species D. melanogaster from the same area. Both desiccation and starvation resistance showed no clinal pattern despite heritable variation among isofemale lines. In contrast chill coma resistance exhibited a linear cline in the anticipated direction, resistance increasing with latitude. Body size was measured as wing length and body weight. Both traits showed geographic variation and strong non-linear clines with a sharp reduction in size in the tropics. These results are discussed in the context of climatic selection and evolutionary processes limiting species borders.
Here, we describe a randomization testing strategy for mapping interacting quantitative trait loci (QTLs). In a forward selection strategy, non-interacting QTLs and simultaneously mapped interacting QTL pairs are added to a total genetic model. Simultaneous mapping of epistatic QTLs increases the power of the mapping strategy by allowing detection of interacting QTL pairs where none of the QTL can be detected by their marginal additive and dominance effects. Randomization testing is used to derive empirical significance thresholds for every model selection step in the procedure. A simulation study was used to evaluate the statistical properties of the proposed randomization tests and for which types of epistasis simultaneous mapping of epistatic QTLs adds power. Least squares regression was used for QTL parameter estimation but any other QTL mapping method can be used. A genetic algorithm was used to search for interacting QTL pairs, which makes the proposed strategy feasible for single processor computers. We believe that this method will facilitate the evaluation of the importance at epistatic interaction among QTLs controlling multifactorial traits and disorders.