Current Anthropology Volume 50, Number 5, October 2009 603

Rethinking the Origins of Agriculture

Health versus Fitness

Competing Themes in the Origins and Spread of Agriculture?

by Patricia M. Lambert

The general picture of human health that has emerged from bioarchaeological studies of the agri-
cultural transition is one of health decline, although the nature and severity of the biological impacts
have varied in accordance with worldwide diversity in the timing, duration, and specific characteristics
of this economic shift. Conversely and somewhat paradoxically, the emerging picture has also been
one of enhanced fertility and population growth. These findings raise challenging questions about
the measures bioarchaeologists use to assess the biological costs and benefits of agriculture. It is
argued here that these measures fall into two potentially quite distinct categories—physiological
fitness (homeostasis) and reproductive (Darwinian) fitness, measures that may assess the costs and
benefits of a biocultural system very differently. Both provide valuable insights into questions about
our past at levels ranging from the evolution of our species to the unique experiences of individuals
and their kin. However, the relative importance of each in larger questions about human adaptation
needs to be carefully considered when assessing the biological evidence in questions of causation.

The role of bioarchaeology in the study of agricultural origins
has been primarily one of biological cost-benefit assessment.
Bioarchaeologists appeal to osteological data, and to a lesser
extent data from other tissues and biological products such
as calculus and coprolites, to reconstruct diet and determine
to what extent a shift in subsistence impacted or was impacted
by the health of individuals and groups. The morphology and
composition of the human skeleton reflect the direct inter-
action of humans with their environment and therefore pro-
vide an empirical basis for evaluating the biological costs and
benefits of different subsistence regimes. Patterning in bio-
logical markers across individuals provides a basis for iden-
tifying shared experiences and thus for evaluating the impact
of different subsistence strategies at the level of the population.
Population-level comparisons in turn provide a basis for gen-
eralizing experiences of populations representing different
economies and subsistence regimes. Bioarchaeological studies
have tended to focus on the cumulative picture of diet, nu-
trition, disease experience, workload, and trauma in popu-
lations before and after the transition from foraging to farm-
ing (Cohen and Armelagos 1984; Cohen and Crane-Kramer
Patricia M. Lambert is Professor in the Anthropology Program at
Utah State University (Old Main 0730, Logan, Utah 84322-0730,
U.S.A. [patricia.lambert@usu.edu]). This paper was submitted 10 IX
08 and accepted 1 V 09.
2007; Larsen 1995, 1997), and conclusions concerning the
biological consequences have correspondingly centered on
changing patterns of health (broadly defined) with the adop-
tion and spread of agriculture. One of the conundrums
emerging from this research is that results of these health-
based studies are often at odds with results of paleodemo-
graphic reconstructions, which suggest enhanced fertility and
population growth with agriculture (e.g., Bocquet-Appel and
Naji 2006; Bocquet-Appel et al. 2005). The purpose of this
paper is to address the meaning of this divergent pattern of
health versus reproductive outcomes at the transition to ag-
riculture by clarifying the two potentially very different bi-
ological measures by which biological anthropologists assess
biological costs and benefits: physiological fitness (homeosta-
sis) and reproductive (Darwinian) fitness (Harrison 1998).
When bioarchaeologists examine human skeletons for var-
ious skeletal markers of health and disease, their goal is to
establish the degree to which individuals and the groups they
comprise were able to maintain homeostasis in different en-
vironments and in the face of changing biocultural systems.
Homeostasis is defined as "a relative constancy in the internal
environment of the body, naturally maintained by adaptive
responses that promote healthy survival" (Anderson and An-
derson 1994, 498). The suite of biological indicators used to
evaluate homeostasis in individuals and groups at agricultural
transitions has grown to include a wide range of skeletal mark-

© 2009 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2009/5005-0003$ 10.00. DOI: 10.1086/605354
 604
ers that in concert permit a general characterization of health:
dental caries, periodontal disease, and antemortem tooth loss
as indicators of dental health; enamel hypoplasia and micro-
defects in tooth enamel, dental asymmetry, tooth size, juvenile
growth patterns, and adult stature as indicators of grovrth
disruption and attainment; porotic hyperostosis as an indi-
cator of iron deficiency anemia and scurvy; periosteal lesions
as indicators of infection and to a lesser extent a number of
other health-related conditions; skeletal profiles for diseases
such as treponematosis and tuberculosis that affect the skel-
eton and convey both disease exposure and risk of infection;
osteoarthritis as an indicator of degenerative joint disease;
and various types of traumatic injury as indicators of behav-
ioral risks associated with different socioeconomic systems
(Lambert 2007; Larsen 1995, 1997; Powell, Bridges, and Mires
1991; Powell and Gook 2005; Roberts and Buikstra 2003).
The general picture that has emerged from such studies is
one of health decline with the transition to and spread of
agriculture (Gohen and Armelagos 1984; Gohen and Grane-
Kramer 2007; Larsen 1995, 1997; Powell, Bridges, and Mires
1991), although workload has often been shown to decline
(Gohen and Armelagos 1984; Larsen 1995, 1997) and vari-
ability across biological indicators attests to the complexity
of interactions between human biological systems and ex-
ogenous environmental systems (e.g., Lambert 2000). Al-
though dietary change is certainly implicated in these chang-
ing patterns of health, especially dental health, concomitant
lifestyle changes such as increasing sedentism, food storage,
clearing of native vegetation, and population growth are also
important correlates of the observed health outcomes (Lam-
bert 2007; Larsen 1995, 1997).
It should be noted that this characterization of health de-
cline based on skeletal health indicators has not gone un-
challenged. In 1992, James Wood and colleagues published a
critique of the osteological evidence brought to bear on the
question of agricultural origins in their seminal paper, "The
Osteological Paradox" (Wood et al. 1992). These authors ar-
gued that bioarchaeologists had been incautious or simply in
error in their interpretations of data from death assemblages
because they failed to recognize or acknowledge that such
assemblages differ in fundamental ways from the living pop-
ulations that produce them. They also pointed out that the
interpretation of skeletal lesions across age categories required
a consideration of both frailty and survivorship and that fail-
ure to do so could lead to erroneous interpretations about
relative health. A case in point was enamel hypoplasia, a lesion
of tooth enamel that forms when growth is disrupted by
systemic disturbance (Goodman, Armelagos, and Rose 1980;
Kreshover 1944, 1960). The authors reasoned that individuals
with these lesions have to survive the stress episode to express
them and that affected individuals thus might more accurately
be viewed as advantaged survivors rather than disadvantaged
victims. They argued that advantaged populations, such as
agriculturalists, might show the lesions more often than dis-
Current Anthropology Volume 50, Number 5, October 2009
advantaged populations, such as foragers, because they sur-
vived long enough to develop them.
These criticisms still resonate in the bioarchaeological lit-
erature on agricultural origins and other human transitions,
although new indicators have been added to the repertoire
and much has been learned about indicators such as enamel
hypoplasia in the intervening years that suggests that the prob-
lems were not as crippling as potentially conceived. For ex-
ample, more recent studies of enamel hypoplasia in living
populations do not support the argument that populations
with a high prevalence of enamel hypoplasia are advantaged
relative to those with a low prevalence (e.g., Guitelli-Steinberg
and Benderlioglu 2006; Lukacs and Walimbe 1998; Zhou and
Gorruccini 1998). Zhou and Gorruccini (1998), for example,
demonstrated that enamel hypoplasia was indeed most prev-
alent in the sector of the Ghinese population (rural) most
impacted by morbidity and mortality during the great famine
of 1959-1961. Although the picture of health at the agricul-
tural transition is more complex than once argued as research
on the topic has broadened to encompass a wide range of
different agricultural products, food processing and storage
techniques, climatic regimes, natural pathogens, and so forth,
health decline continues to predominate as the more common
experience for emerging agriculturalists. This is seen in in-
creasing evidence for dental disease, growth disruption, in-
fectious disease, and nutritional deficiency (e.g., Gohen and
Grane-Kramer 2007; Lambert 2000; Larsen 1995, 1997).
The question might reasonably be asked as to why people
in so many world regions adopted an economic strategy that
almost universally resulted in declining health. A number of
scholars over the years have argued that such economic in-
tensification was born of necessity: populations grew, natural
resources were insufficient to feed the growing population,
and population pressure necessitated resource intensification
(e.g., Boserup 1965; Gohen 1977; Earle 1980; Winterhaider
1981). This could certainly be true in the case of agriculture,
though the actual transition has been more difficult to eval-
uate biologically than the health of forager antecedents relative
to their agriculturalist descendants, thus hampering efforts to
establish a causal link. However, it is also possible that the
primary measure used to assess the biological costs of the
agricultural transition has missed one fundamental compo-
nent of the cost-benefit equation: reproductive fitness.
Beginning in the 1980s paleodemographers and other re-
searchers involved in the study of ancient populations began
to challenge results of life table analysis based on data from
prehistoric skeletal series as indicative of increased mortality
and reduced life expectancy with the transition to agriculture.
They pointed out that death assemblages were not necessarily
reflective of living populations and that changes in fertility
rather than changing mortality likely accounted for apparent
reductions in life expectancy (Hershkovitz and Gopher 1990;
Johansson and Horowitz 1986; Sattenspiel and Harpending
1983; Walker, Johnson, and Lambert 1988; Wood et al. 1992).
In consequence, innovative new methods were developed for
 Lamben Health versus Fitness
assessing reproductive rate and population growth in the past.
The emerging picture is one of increasing fertility and pop-
ulation growth with the shift to agriculture in both Old and
New World contexts (e.g., Bocquet-Appel 2007; Bocquet-
Appel and Naji 2006; Bocquet-Appel et al. 2005), with the
initial shift to cultigens and later to Old World domesticated
animals argued by some to have been of particular importance
in the New World (McCaa 2002). In other words, in terms
of reproductive outcomes, agriculture appears to have been
life enhancing.
The two measures of the biological costs associated with
agriculture thus produce seemingly conflicting results: health
decline and enhanced reproduction. These apparently incom-
patible findings beg validation and explanation from a mod-
ern context where the relationship between health and re-
productive rate can be accurately assessed without concerns
about sampling biases and other potential problems that com-
plicate the interpretation of death assemblages. For several
reasons modern Africa provides an ideal context for such a
study. First, agricultural production is the primary focus of
the African economy, employing more than half of the African
labor force (Adewumi 2008; Barrios, Ouattara, and Strobl
2008; table 1, percent rural), and is therefore reflective of the
mode of production of interest in this study. Second, many
African countries continue to struggle with malnutrition
(United Nations 2006) and high rates of infectious diseases
such as cholera and HIV/AIDS (Gaffga, Tauxe, and Mintz
2007; Gregson et al. 2007; Mugoya et al. 2008; Shears 2007),
making Africa a logical choice for investigating the impacts
of malnutrition and infectious disease on fertility and growth
rate. Third, demographic data are readily available for most
African countries (United Nations 2006; UN Statistics Divi-
sion 2005, 2007(3, 2007Í;, 2007c), making it possible to explore
this relationship with a statistically valid data set. The Africa
data can provide a basis for interpreting what the bioarchaeo-
logicai and paleodemographic observations described above
might mean in terms of the viability of individuals, popula-
tions, and economic strategies in the past.
Health and Reproductive Fitness in
Modern Africa
In order to explore the relationship between health and re-
productive rate, data on undernourishment in 46 African
countries were obtained online from the UN World Food
Bank (United Nations 2006; table 1). These data were used
as a proxy measure of health risk: the greater the percent of
undernourishment, the greater the assumed risk of dimin-
ished health. Data on fertility, annual growth rate, per capita
GDP, and urban/rural location for 51 African countries were
also obtained online from the UN Statistics Division (2005,
2007fl, 2007b, 2007c). These data provide a basis for statis-
tically evaluating the nature of the relationship between phys-
iological homeostasis and reproductive fitness.
Analysis of correlation reveals a significant, positive cor-
605
relation between undernourishment and fertility (Spearman's
p = 0.459, p = 0.001) in these African countries: the most
malnourished people tend to have the highest fertility. Fur-
ther, fertility is negatively correlated with per capita GDP
(Pearson's r = -0.360, p = 0.009), indicating that the pop-
ulations with the lowest incomes tend to have the highest
birthrate (see also Dow et al. 1994). Of course, fertility rate
is not reproductive rate, and it could be argued that birthrate
is not indicative of growth rate. However, correlation analysis
of these data also reveals a strong positive correlation between
fertility and growth rate (Pearson's r = 0.724, p<0.001), so
despite poor nutrition, high levels of infectious disease, and
relatively high infant mortality (UN Statistics Division 2008),
populations are growing. In other words, homeostasis as mea-
sured by health and nutrition is compromised without greatly
affecting fertility and population growth—at least for a time.
Clearly, HIV has impacted population growth in some African
countries (Gregson et al. 2007), and it is likely that disease
epidemics also emerged among settled farmers in prehistory
that impacted growth and may account for changing profiles
and population collapses seen later in agricultural sequences
(e.g., Bocquet-Appel 2007; Gohen and Grane-Kramer 2007).
Discussion and Conclusions
The point of this analysis is not to explore why fertility is
highest in many of these undernourished, largely rural pop-
ulations, although the causes of fertility shifts have figured
centrally in discussions of population growth in Africa (e.g.,
Boserup 1985; Dow et al. 1994) and in prehistoric populations
at the origins of agriculture (e.g., Gashdan 1985; Handwerker
1983, 1985; Knauft 1987; Nardi 1981; Roth 1985, 1993).
Rather, these data are presented because they illustrate the
counterintuitive finding that health can be substantially com-
promised by disease and poor nutrition, with little impact on
reproductive rate, and that this could be what is seen world-
wide in human populations transitioning to agriculture. The
significance of these findings in terms of questions concerning
the origins and spread of agriculture is that while the health
changes seen with this economic transition are profoundly
important for documenting the biological costs of agriculture
and the agricultural lifestyle, they may be less informative
about the causes of this transition. This is not to say that
population pressure played no role as a "push" mechanism,
particularly in the origins of agriculture (as per Boserup 1965;
Gohen 1977; Earle 1980; Winterhaider 1981), but rather that
the biological data also support an alternative model for its
adoption and spread. Instead of declining health and life qual-
ity as a prime mover (or proxy measure of a population
pressure model) for this economic transition, it might have
come about in consequence of a biological "pull" mechanism:
increased fertility and population growth rate. Indeed, the
data at hand suggest that agriculture may have emerged and
spread because it was a behavioral strategy that enhanced
reproduction. As demonstrated above, health could have de-
Table 1. Recent demographic statistics for Africa

Per capita Percent Fertility Annual Percent

GDP undernourished rate growth rate rural
African country 2006' 2006'' 2005' 2000-2005'' 2005'

Algeria 3,476 5-19 2.5 1.5 36.7

Angola 2,855 35+ 6.8 2.8 46.7
Benin 536 5-19 5.9 3.2 59.9
Botswana 4,755 35-t- 3.2 .1 42.6
Burkina Faso 416 5-19 6.7 3.2 81.7
Burundi 114 35+ 6.8 3.0 90.0
Cameroon 1019 20-34 4.6 1.9 45.4
Cape Verde 2,153 3.8 2.4 42.7
CAR 333 35+ 5.0 1.3 62.0
Chad 634 20-34 6.7 3.4 74.7
Comoros 486 4.9 2.6 63.0
Congo 1,946 35+ 6.3 3.0 39.8
Cote d'Ivoire 952 5-19 5.1 1.6 55.0
Djibouti 925 5.1 2.1 13.9
DRC 136 35+ 6.7 2.8 67.9
Egypt 1,484 2.5-4 3.3 1.9 57.2
Equatorial Guinea 19,166 20.34 5.9 2.3 61.1
Eritrea 249 35 + 5.5 4.3 80.6
Ethiopia 164 35 + 5.9 2.4 84.0
Cabon 7,245 5-19 4.0 1.7 16.4
Gambia 307 20-34 4.7 2.8 46.1
Ghana 532 5-19 4.4 2.1 52.2
Guinea 311 20-34 5.9 2.2 67.0
Kenya 650 20-34 5.0 2.2 79.3
Lesotho 725 5-19 3.6 .1 81.3
Liberia 192 35 + 6.8 1.4 41.9
Libya 8,348 2.5 3.0 2.0 15.2
Madagascar 287 35+ 5.4 2.8 73.2
Malawi 164 35 + 6.1 2.3 82.8
Mali 498 20-34 6.9 3.0 69.5
Mauritania 899 5-19 5.8 3.0 59.6
Mauritius 5,124 5-19 2.0 1.0 57.6
Morocco 2,087 5-19 2.8 1.5 41.3
Mozambique 349 35 + 5.5 2.0 65.5
Namibia 3,084 20-34 4.0 1.4 64.9
Niger 247 20-34 7.9 3.4 83.2
Nigeria 917 5-19 5.8 2.2 51.8
Reunion 2.5 1.6 7.6
Rwanda 242 35 + 5.7 2.4 80.7
Senegal 768 20-34 5.0 2.4 58.4
Seychelles 8,209 2 .9 47.1
Sierra Leone 318 35 + 6.5 4.1 59.3
Somalia 283 35 + 6.4 3.2 64.8
South Africa 5,133 2.8 .8 40.7
Sudan 934 20-34 4.4 1.9 59.2
Swaziland 2,399 5-19 4.0 .2 75.9
Togo 356 20-34 5.4 2.7 59.9
Tunisia 3,003 2.5 2.0 1.1 34.7
Uganda 346 5-19 7.1 3.4 87.4
U. R. Tanzania 335 35 + 5.0 2.0 75.8
Zambia 938 2.5 5.7 1.7 65.0
Zimbabwe 133 35+ 3.6 64.1

'UN Statistics Division 2007b.

•"United Nations 2006.
'UN Statistics Division 2005.
"'UN Statistics Division 2007c.
'UN Statistics Division 2007fl.
 Lamben Health versus Fitness
dined substantially, but so long as it did not impact repro-
ductive rate, populations practicing agriculture would have
differently expanded.
A Darwinian model thus might better account for the pace
and scale of the agricultural transition than one based on
health and physiological homeostasis. Health decline could
have been a costly by-product of an economic system that
enhanced fertility and led to population growth. As long as
it did not affect reproductive rate, declining health may have
wielded little influence on how people chose to make a living.
People would likely have been unaware of the association
between health decline and agriculture, at least in terms of
problems such as bacterial infection and its link to dental
caries and other infectious diseases. Changes in fertility could
have come about in consequence of conscious decisions about
age at marriage, birth spacing, contraception, and/or other
forms of birth control (e.g., Cashdan 1985; Harris and Ross
1987), but it is equally plausible that physiological changes
in fertility (and fecundity) associated with declining mobility,
more regular food supply, and decreased workload accounted
for this demographic transition (e.g., Handwerker 1983; Roth
1985). It is therefore conceivable that one of the greatest
human cultural transitions was driven by the same evolu-
tionary forces responsible for so many other important de-
velopments in the history of our species: a positive feedback
relationship between cultural behavior and reproductive
fitness.
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