AMERICAN JOURNAL OF HUMAN BIOLOGY 18:1–9 (2006)

Pearl Memorial Lecture

Biocultural Approaches in Human Biology
DARNA L. DUFOUR*
Department of Anthropology, University of Colorado, Boulder, Colorado 80309-0233

ABSTRACT Biocultural approaches recognize the pervasiveness and dynamism of interac-

tions between biological and cultural phenomena, and they explicitly strive to integrate biological,
sociocultural, environmental, and other kinds of data. They have been part of human biology at
least since 1958, when Frank Livingstone so elegantly explained the linkages among population
growth, subsistence strategy, and the distribution of the sickle cell gene in West Africa. These
approaches developed further with the advent of human adaptability studies in the 1960s as part
of the Human Biological Program and have become increasingly focused on understanding the
impacts of everyday life on human biological variation. Biocultural approaches generate explana-
tions that are intuitively appealing to many because they offer a kind of holistic view. They can,
however, be very challenging approaches to implement, perhaps in part because we are more
experienced in measuring the biological than the cultural. Some of the challenges include (1)
defining precisely what we mean by constructs like socioeconomic status, poverty, rural, and
urban; (2) operationalizing key variables so that they can be measured in ways that are ethno-
graphically valid as well as replicable; (3) defining and measuring multiple causal pathways. In
this paper, I briefly review the history of biocultural approaches and then illustrate some of the
challenges that these approaches present with examples from my own research on nutrition and
energetics as well as that of other practitioners. Am. J. Hum. Biol. 18:1–9, 2006. # 2005 Wiley-Liss, Inc.

Human biology is an interdisciplinary able as well as ethnographically valid; and (3)

field centrally concerned with understand- defining and measuring multiple causal
ing human biological variability and the pathways. In this paper, I briefly review the
mechanisms responsible for that variability. history of biocultural approaches and then
Researchers represent a variety of perspecti- illustrate some of the challenges that these
ves, particularly those that are comparative, approaches present with examples from my
developmental, ecological, and/or evolution- own research on nutrition and energetics.
ary. Within each of these perspectives, bio-
cultural approaches are those that explicitly
recognize the dynamic interactions between
humans as biological beings and the social, A BRIEF HISTORY
cultural, and physical environments they
It is difficult to know where to begin, but
inhabit. They focus on understanding varia-
because this is the Pearl Memorial Lecture,
bility in human biology as a function of
it is fitting that we start with Raymond
responsiveness to the larger (social, cultural,
Pearl. Although Pearl is best known for his
and physical) environment.
contributions to biostatistics, he clearly
Biocultural approaches have a long history
understood the impact of the social and cul-
in human biology and biological anthropol-
tural environment on human biology and
ogy, a closely linked scientific community.
employed culturally defined variables in
They can, however, be very challenging app-
many of his analyses. The importance Pearl
roaches to implement, perhaps in part
attributed to the social and cultural environ-
because we are more experienced in measur-
ment is clear in the following passage:
ing the biological than the social or cultural.
Some of the challenges include (1) defining
precisely what we mean by constructs like *Correspondence to: Darna L. Dufour. E-mail: Darna.
Dufour@colorado.edu
socioeconomic status, poverty, social support, Received 29 August 2005; Accepted 19 September 2005
etc.; (2) operationalizing key variables so that Published online in Wiley InterScience (www.interscience.
they can be measured in ways that are replic- wiley.com). DOI: 10.1002/ajhb.20463

ß 2005 Wiley-Liss, Inc.

2 D.L. DUFOUR
‘‘ . . . economic and social factors and forces
are among the most important elements in
determining the biologically significant
environment of human beings, as they exist
here and now. Relative wealth virtually
determines the character of the physical
environment in which men live.’’ (Pearl,
1930:540)
Pearl was ahead of his time in considering
the impact of the social and cultural environ-
ment on human biology, at least in com-
parison to the discipline of anthropology;
physical anthropologists were focused on
issues of race, and their work reflected the
extreme hereditarianism of the time. The
dominant paradigm of physical anthropology
began to shift away from issues of race in the
1940s and 1950s to one more focused on evo-
lution and natural selection and on attempts
to understand human biological variation as a
result of adaptation to the environment. The
environment of interest was principally
the physical environment, but the paradigm
shift opened the door to the consideration
of ‘‘environment’’ more broadly defined.
One of the first people to clearly conceptua-
lize the ‘‘environment’’ as more than the
external physical conditions surrounding a
human population was the late Frank
Livingstone, and his famous 1958 paper on
malaria and the sickle cell trait in Africa
was a landmark in biocultural approaches.
The paper is probably best remembered as
the model of disease as an agent of natural
selection; the disease being malaria, and the
adaptation being genetic, i.e., the sickle cell
trait. But the paper is much more than that.
It is really an elegant explanation of the lin-
kages among population growth, subsistence
strategy, the natural history of the mosquito
as a disease vector, and the distribution of the
sickle cell gene in West Africa. It is a brilliant
analysis, and certainly a biocultural one.
Livingstone’s contribution notwithstand-
ing, the dominant idea remained, however,
to consider the environment as the physical
environment. This is clear in the first text-
book in human biology, the 1964 text Human
Biology—An Introduction to Human Evolu-
tion, Variation and Growth by Harrison et
al. The text has a section entitled ‘‘Human
Ecology’’ in which the late J.S. Weiner defines
the environment as follows:
The ‘‘environment’’ represents the totality
of the surroundings in which the community
finds itself, and includes physical and living
elements, that is, the geology, topography,
and climate of the terrain and its communi-
cations; the vegetational cover and the insect
animal and bird life . . . The natural environ-
ment may be partly or largely replaced by a
domesticated or industrialized environment
of shelters, gardens and farmlands or even
by complete urbanization. (Weiner, 1964:401)
The chapters in the section, ‘‘Nutritional
Ecology,’’ ‘‘Climate Adaptation,’’ ‘‘Disease
and Population Stability,’’ were all very bio-
logically oriented and showed little concern
with variables that might be considered part
of the larger cultural or social environment.
This orientation to the environment as lim-
ited to the physical environment character-
ized the early stages of the human adap-
tability paradigm.
If we assume, with Kuhn (1970), that text-
books articulate the paradigms to which the
scientific community is committed at a given
point in time, we can use them to trace the
development of biocultural approaches in
human biology and biological anthropology.
The next two human biology textbooks,
published in the 1970s, demonstrated more
of an interest in incorporating the role of the
social and cultural environment into our
understanding of human biological variation.
The first, Ecology, Energetics and Human
Variability, by Little and Morren (1976), was
a collaboration between a biological and a cul-
tural anthropologist. It covered much of the
same material in human ecology as did the
1964 textbook, but it also included a series of
case studies of traditional populations in dif-
ferent ecosystems that attempted to integrate
the work done by biological anthropologists
with that of ecologically oriented cultural
anthropologists. The case-study approach
was hugely influential because it demon-
strated the potential of a more holistic view
of human populations. It probably served as
the template for Moran’s (1979) widely used
textbook in ecological anthropology, Human
Adaptability. The second textbook, Human
Adaptation: A Functional Int-erpretation by
Frisancho (1979), was more narrowly focused
on biological, especially physiological,
responses to stresses of the physical environ-
ment. It did, however, include a chapter on
the westernization of dietary habits and dis-
ease expression that has a biocultural focus.
It was around the same time that the term
‘‘biocultural’’ started to appear in the litera-
ture. The first paper in the journal Human
 BIOCULTURAL APPROACHES IN HUMAN BIOLOGY
Biology with the term ‘‘biocultural’’ in the
title was ‘‘Biocultural correlates of blood pres-
sure of Samoan migrants in Hawaii,’’ by
Hanna and Baker (1979). The ‘‘biocultural’’
correlate was place of residence, namely, rural
versus urban.
The third and last edition of the Harrison et
al. textbook, Human Biology, was published
in 1988. In that edition, Baker replaced
Weiner and renamed the section on human
ecology to ‘‘Human Adaptability.’’ The chap-
ters remained essentially the same, except
that the chapter on population stability was
replaced by one entitled ‘‘Human Biological
Responses to Modernization.’’ This chapter
reflected the ongoing research in Samoa by
Baker and colleagues that had the explicit
goal of understanding the effects of moderni-
zation, a culturally defined construct, on biol-
ogy (Baker 1986). The Samoan research
exemplified the deepening interest among
biological anthropologists in culturally
defined independent variables and a willing-
ness to consider the social and cultural com-
ponents of the environment.The most recent
textbook, Human Biology: An Evolutionary
and Biocultural Perspective, edited by
Stinson et al. (2000) was written collabora-
tively by members of the Human Biology
Association. The book includes the term ‘‘bio-
cultural’’ in the title and in the Preface; the
editors state:Human biology relies heavily on
an evolutionary perspective to explain varia-
tion through space and time but also consid-
ers to be crucial the effect that human
cultures have on our biology—a biocultural
perspective. (Stinson et al., 2000)The chap-
ters cover what are currently considered to
be the major areas within human biology:
genetic variation, variation due to climate,
disease, nutrition and energetics, growth and
aging, and demography. Most of the chapters
employ a concept of the environment that
includes social and cultural components as
well as physical ones, and hence they embrace
a biocultural perspective. This textbook then
brings us to the present day in which biocul-
tural approaches play a central role on human
biology and biological anthropology.
Beginning in the mid- to late-1990s there
has been a call within biological anthropology
to broaden the scope of biocultural analyses
by including insights from political economy.
This idea is most forcefully articulated in
the volume, Building a New Biocultural
Synthesis, edited by Goodman and Leather-
man (1998). In particular, the editors place a
3
more intensified focus on the social, political,
and economic forces that affect health. The
rationale here is basically that the biological
condition of the subjects we wish to under-
stand is responsive to and hence affected by
the social and cultural environment. So, in
some ways we have come full circle, and are
cognizant of what Raymond Pearl pointed out
in 1930, i.e., that ‘‘ . . . economic and social
factors and forces are among the most impor-
tant elements in determining the biologically
significant environment of human beings . . . ’’
(Pearl, 1930:540).
CHALLENGES
Biocultural approaches can be very chal-
lenging approaches to implement. I have
struggled with these challenges in my own
work, and I am indebted to both McElroy
(1990) and Dressler (1995) for helping me
think about some of the challenges in a
more formal way. Here, I would like to focus
on my attempts to deal with three the types
of challenges. They are: (1) understanding
the meaning of a major construct like
poverty; (2) operationalizing key vari-
ables so that they can be measured in ways
that are ethnographically valid as well as
replicable; and (3) defining and measuring
multiple causal pathways.
Understanding the meaning of constructs
like poverty
‘‘Poverty’’ is a major construct I have dealt
with in my own work although not always as
thoughtfully as I might have done. What
does it really mean to say people are ‘‘poor’’
or live ‘‘in poverty,’’ or, even closer to home,
what do we mean by the ‘‘biology of
poverty’’? Dictionary definitions of poverty,
like, ‘‘having little or no money and few or
no material possessions’’ (Wordnet, 2003),
are not very helpful because poverty is really
more than that. As Narayan (2000) has
argued, poverty is a multidimensional social
and economic phenomenon, and definitions
can vary by social and economic context.
Although it is typically defined as a lack of
material resources needed to maintain well-
being, food being pre-eminent among these
resources, other material assets like housing
and land can also be important. Probably
less recognized is the fact that poverty also
has an important psychological component
that is related to feelings of powerlessness,
4 D.L. DUFOUR
social exclusion, humiliation, and depen-
dency (Narayan, 2000).
I have been particularly interested in the
relationship between poverty and nutrition,
and I would like to discuss it here in the
context of one research project, The Cali
Project. This was a project developed by G.B.
Spurr in the later 1980s; I was a co-primary
investigator along with Julio C. Reina, a
medical doctor. The aim was to understand
the impact of undernutrition on the biology
and behavior of women living in poverty in
Cali, Colombia. The initial model was an
adaptive-type model, with undernutrition
being the stressor and the responses of
interest being biological and behavioral
accommodations to undernutrition. The
guiding assumption, based on the literature
available at the time, was that in a develop-
ing country like Colombia, poverty would be
major determinant of undernutrition. To
find women living in poverty we focused on
the sectors of the city of Cali defined as very
low SES (socioeconomic status), and specifi-
cally on two neighborhoods, or barrios: a
squatter settlement and a planned, but
illegal, urbanization.
In the beginning of the project, I thought I
knew what poverty was; that is, I thought I
would know it when I saw it. But my concept
of poverty was vague. It was useful in broad
comparisons of rich versus poor but too
fuzzy to help us understand the impact of
the social and economic condition known as
‘‘poverty’’ on people’s biology.
The first thing we did in the project was an
anthropometric survey to define the general
level of nutritional status in the city of Cali.
We surveyed three SES groups (a low, a
mid–low, and a high) following the defini-
tions provided by the municipal government
(Dufour et al., 1994). SES differences in
stature were where we expected them to be;
the high-SES women on about the U.S. 25th
percentile and the lower-SES women on
about the 10th percentile, suggesting under-
nutrition during growth and development.
SES differences in BMI, were not, however,
what we expected to find. First, there were
very few women (3–5%) we could classify as
undernourished based on BMI (i.e., a BMI of
18.5 or less). Second, the mean BMI in the
low-SES group was greater than in the high-
SES group. Third, twice as many of the low-
SES women were overweight (based on the
standard cut off of a BMI > 25 (36% of low-
SES women as compared to 16% of high-SES
women). The greater prevalence of over-
weight in the low-SES women conformed to
a pattern seen in industrialized countries
like the U.S. but is just the opposite of what
we had expected to find in a developing
country like Colombia in the early 1990s.
Even though our measure of SES lacked
precision, the higher mean BMI in the low-
SES group was a question begging for a
biocultural approach—one with real atten-
tion to the cultural component.The ap-
proach I took was an inductive one: to try
to understand the links between poverty and
nutritional status in this particular ethno-
graphic context. As a start, I spent some
time doing ethnographic observations and
informal interviews. Those initial ethno-
graphic observations and informal interviews
proved to be invaluable and suggested that
the nutritional situation was not a rosy as
the BMI data suggested. I learned that:
* In the poorest households there was no
food storage—not even of things like salt
and sugar; food was purchased on a meal-
to-meal basis. In other households, staples
like rice, sugar, and coffee were purchased
weekly, and everything else on a daily
basis.
* The main meal of the day was the mid-day
meal and typically consisted of rice, a
small portion of beef (about 1 oz.), a bit of
tomato and maybe onion, and a sugared
drink. The morning and evening meals
were smaller and sometimes quite mini-
mal, like only sugared coffee.
* One woman said she had not had much to
eat for two days because her family had
had to pay a medical bill, and hence did
not have much money for food.
* Some preschool children showed evidence
of resting postures associated with energy
deficits; some adults also showed evidence
of the same.
* The inclusion of beef in the mid-day meal
was important culturally, but for some
people the ‘‘cuts’’ of meat were unusual:
cow’s hoof, ‘‘cheek’’ meat, esophagus.
These were cuts were considered unusual
not only by us, but also by the women who
lived in the barrios, and were evidence of
not enough money to buy more acceptable
forms of animal protein. This behavior did
not necessarily change the nutritional
value of the diet, but it did lead to humilia-
 BIOCULTURAL APPROACHES IN HUMAN BIOLOGY
tion because the foods were socially inap-
propriate.
As the study progressed over the next 4
years or so, these initial impressions were
confirmed by further observations and inter-
views.
The diet records we collected also provided
evidence of days people did not eat very
much (Dufour et al., 1997). We defined these
as ‘‘low energy intake’’ days, i.e., days when
energy intake was less than or equal to 1.27
times BMR. This value is what the FAO/-
WHO/UNU (1985) calls the ‘‘survival’’ re-
quirement; it is about equivalent to the
energy needs of a bed-ridden adult. Of the
nonpregnant, nonlactating women in the
study, 41% of them had 2–6 low-intake days
out of 6 days total in approximately 6
months. We did not investigate the social
and economic circumstances surrounding all
of these low-intake days, but we did rule out
illness as a possible cause as well as ‘‘diet-
ing’’ (a very rare behavior in this popula-
tion). Hence, we assumed that these ‘‘low-
intake’’ days represented economic con-
straint to food acquisition. Because average
BMI was normal or above, we also assumed
that these low-intake days were offset by
higher-intake days, so that average energy
intake was adequate over the long run. So,
in the case of the Cali women, what does
poverty mean in this one dimension of
nutrition? It means short-term fluctuations
in food intake that reflect fluctuations in
money that can be used to buy food. ‘‘Short-
term’’ here is day-to-day for most households,
week-to-week for others. Some of the fluctua-
tions are predictable, most are not. There are
no seasonal effects. As one woman said,
‘‘When you have money you eat a lot, and
when you don’t have money you don’t eat.’’
Does this help us understand the impact of
poverty on biology? I think so. In comparison
to women with only one or less low-intake
days, those with 2–6 low-intake days had
significantly lower energy intakes (2273 ver-
sus 1833 kcal/day) and activity levels (total
daily energy expenditure (TDEE)/BMR ¼ 1.95
versus 1.71), and a greater percentage of them
had inadequate protein and micronutrient
intakes (Dufour et al., in preparation). BMI
was similar in the two groups and hence was
not very informative. There are at least two
reasons for this. First, BMI measured at any
one point in time is the result of a cumulative
history of energy balance. In the Cali women,
5
energy balance was apparently more positive
than negative over the long term, and hence
BMI did not capture the relationship between
poverty and diet in this group. Second, BMI is
a very coarse measure of nutritional status
because the ‘‘normal range’’ (18.5–25 kg/m) is
very wide: for a Cali woman of average
stature, the average is about 17 kg. Hence,
in an environment like Cali where the food
supply fluctuates over the short term, BMI
does not tell us much.The nutritional dimen-
sion of poverty in the Cali case is different
than poverty in the case where food energy
intake is chronically deficient and BMI very
low, as well as the case where food intake is
deficient seasonally, and BMI shows seasonal
fluctuations. It is also different than the case
of poverty in some parts of the U.S., where
food energy intake is chronically high. So, in
the dimension of nutrition, the meaning of
poverty is context specific. In that regard, the
phrase ‘‘biology of poverty’’ is misleading
because it suggests that poverty is the same
everywhere and implies that there is a single
biological response to poverty. I doubt that
that will prove to be the case because poverty
is a multidimensional phenomenon, and the
specifics of the context (ethnographic, demo-
graphic, epidemiological, nutritional, etc., as
well as characteristics of the human-built and
physical environments) should lead to differ-
ences of interest. Hence, we should be think-
ing in terms of ‘‘biologies of poverty,’’ i.e., of
the variability in responses rather than in
terms of a biology of poverty.
Operationalizing variables
We have a well-developed tool kit of bio-
logical measures and are experienced in
measuring biological outcome variables like
weight, height, serum cortisol, etc. These are
dependent variables that we can easily
quantify and operationalize. In studies using
a biocultural approach, researchers typically
want to assess the impact of a culturally
defined variable on some aspect of biology,
i.e., use a culturally defined variable as an
independent variable. These kinds of inde-
pendent variables are typically a challenge
to operationalize. This is especially true if the
independent variable we want to use is a
complex function of interacting variables.
For example: the variable psychosocial stress
could be a function of social support, food
insecurity, and/or the threat of violence. In
6 D.L. DUFOUR
the early days of human adaptability studies,
the independent variables (characteristics of
the physical environment like heat, cold, and
altitude) were easier to measure.
To adequately operationalize social and
cultural variables requires a reasonable eth-
nographic understanding of the local setting.
This is a point Dressler (1995) has empha-
sized. The survey-type interview data we col-
lect does not really go deep enough to give us
the kind understanding of the local ethno-
graphic situation we need. Let me present
one other example from the Cali project. In
that study we used structured interviews to
obtain sociodemographic information, and
information on work status. With regard to
the latter, our plan had been to use work as a
dichotomous variable and compare the en-
ergetics of working versus not-working wo-
men on the assumption that working women
would be under greater energetic stress. So in
the interview we asked two, seemingly
straight forward, questions: (1) Do you work?
(2) What kind of work do you do?
As the study progressed and our under-
standing of the ethnographic situation
improved, we learned that the Cali women
had a conception of ‘‘work’’ that differed from
our own, even though we both used exactly
the same vocabulary to discuss it (Dufour et
al., 2003). We thought of ‘‘work’’ as an
income-earning activity, and also one that
tended to be a regular, steady kind of
employment. The Cali women also thought
of ‘‘work’’ as an income-earning activity but
not necessarily anything like regular steady
employment. We were thinking in terms of a
formal economy, and they worked in a very
informal one. They considered themselves to
be women who ‘‘worked’’ if they engaged in
any activity that provided a monetary in-
come, no matter the time commitment per
day, or whether it was short or long term,
occasional, steady, or even highly unpredict-
able. For example, some women felt that they
‘‘worked’’ for the research project because
they were remunerated for their participa-
tion as subjects. Other women felt they
‘‘worked’’ for the chicken-processing plant,
even though they might only be called in to
work a couple of days a month, and not
necessarily every month. One woman who
said she ‘‘worked’’ spent less than 30 minutes
a day selling a homemade sugared drink
through a window of her home.
The second question (‘‘What kind of work do
you do?’’) was difficult for some women to
answer because their ‘‘work’’ encompassed a
variety of different income-earning strategies,
and their engagement in ‘‘work’’ was highly
opportunistic both in terms of strategy and
time. For some women, the type of ‘‘work’’ they
did varied from day to day or was different
between weekdays and weekends. For exam-
ple, a woman might sell food as a street vendor
on Saturday evening, take in some ironing a
couple of days a week, and maybe help someone
tend a shop for an hour or two a week.
The lesson we learned was that, in order
to use a culturally defined variable, like ‘‘work’’,
one has to understand the local ethnographic
context, i.e., be able to interpret the meaning of
word within the cultural context, not only
speak the language. Geertz’s (1973) notion of
thick description applies here; we need a
‘‘thick’’ understanding. Without it, we end up
doing a remote-sensing kind of fieldwork with-
out the ground truth component.
Defining and measuring multiple causal
pathways
Defining and measuring multiple causal
pathways can be a challenge. Many study
designs are based on examination of the
effect of a single independent variable on an
outcome variable. However, if we are inter-
ested in understanding complex interactions
between biology and culture, then the con-
sideration of multiple variables is invalu-
able. As an example, I would like to discuss a
question I have worked on for a number of
years: Why do Tukanoan Indians in the
northwestern Amazon show a strong pre-
ference for manioc (cassava; Manihot escu-
lenta Crantz) varieties that are toxic and
require extensive processing when they also
cultivate nontoxic varieties? The toxicity of
manioc is due to the presence of cyanogenic
glucosides that are hydrolyzed to hydrogen
cyanide (HCN) when the plant tissue is
damaged. The Tukanoan preference is for
manioc cultivars with high cyanogenic po-
tential (high-CNP), i.e., those referred to as
‘‘bitter.’’ This preference is of interest for at
least two reasons. One is that the preference
for high-CNP over low-CNP (i.e., those
referred to as ‘‘sweet’’), which is common
in many parts of the world where both types
are grown, appears to be an exception to the
general rule that humans select the less
toxic varieties of a given food plant (Johns,
 BIOCULTURAL APPROACHES IN HUMAN BIOLOGY
1990). The other is that the manioc cultivars
from Tukanoan Indians at Yapu (the group
referred to here and the only one for which
we have quantitative data) a very high-CNP,
actually higher than any other group of
cultivars reported in the literature (Dufour,
1988).
Tukanoans recognized that high-CNP cul-
tivars were toxic and correctly associated the
toxicity with a bitter taste and an acrid smell
(Dufour, 1988). Nonetheless, these high-
CNP cultivars were the dietary staple and
provided over 80% of the food energy
(Dufour, 1983). Low-CNP cultivars were a
supplementary food and played a very minor
role in the diet. One of the explicitly stated
objectives of processing and cooking was to
eliminate toxicity and bitterness of taste. We
were able to demonstrate that traditional
processing and cooking techniques actually
eliminated over 90% of the total cyanogens
in fresh roots (Dufour, 1989) and, hence,
were highly adaptive. Nonetheless, because
of the high consumption of manioc-based
foods, cyanogen intake was also relatively
high (Dufour, 1995). This high intake did
not appear to have negative effects on
biology (at least in adults), probably because
consumption was spread out over the course
of the day and nutritional intake was
generally adequate, especially in terms of
the substrates necessary for metabolic de-
toxification (primarily sulfur-containing
amino acids) (Dufour, 1995).
So, the Tukanoan case is one in which
people have increased the risk of exposure to
toxic cyanide compounds in the environment
by their choice of staple crop but then
ameliorate the risk behaviorally (using
time-intensive processing and cooking tech-
niques) and culturally (considering the food-
ways and temporal patterns of food intake).
The question of interest then becomes what
factors influence food crop choice and hence
the potential exposure to toxicity?
I began by asking people why they culti-
vated primarily the high-CNP varieties,
which they referred to as kii. Tukanoan
women each explained it in terms of tradi-
tion: their mother did it that way, and their
mother’s mother before her. Men typically
explained it in terms of their origin myth:
the first woman planted seven varieties of
kii and used the roots to make casabe
(manioc bread, one of the staple foods).
Low-CNP varieties, the men reported, were
7
introduced by rubber gatherers from Brazil
around the 1940s. These explanations were
intuitively appealing because the impor-
tance of history and tradition in a group
such as this cannot be denied. There was,
however, one intriguing incongruity: low-
CNP manioc was introduced at about the
same time as a group of yellow-fleshed high-
CNP cultivars (traditional cultivars were
white-fleshed), and the latter had become
important in the diet whereas the former
had not. The yellow-fleshed cultivars were
used to make fariña (manioc meal), which
was introduced at the same time. So tradi-
tion and history, while important, seemed to
be only part of the answer.
The literature offered other explanations
for the preference for high-CNP manioc in
the Amazon Basin and many parts of Africa,
where the phenomenon also occurs. These
explanations, in approximate order of cur-
rent popularity, are that high-CNP cultivars
(1) are higher yielding (Bruijn, 1983; Cock,
1985); (2) are more resistant to predation
and disease because the cyanogenic gluco-
sides function as a deterrent (Bellotti and
Arias, 1993; Kakes, 1990; Pollard, 1992); (3)
have a higher starch content (Lathrap, 1973;
Sauer, 1950); and (4) are more suitable for
certain foods (Lathrap, 1973). We were able
to find little empirical evidence in the litera-
ture in support of any of these explanations,
but we assumed that they were all worth
testing with the Tukanoans at Yapu.
First, to examine differences in yield, we
(Wilson and Dufour) collected data for
representative gardens in 1986 and 1994.
The mean yield (kg roots/plant) of high-CNP
cultivars tended to exceed that of low-CNP
and the differences were statistically signifi-
cant in the 1994 data (Dufour, 1993; Wilson
and Dufour, 1996, 2002).Second, in terms of
starch content, our results were contra-
dictory: starch content was lower in the
high-CNP cultivars sampled in 1986 but
higher in the 1994 samples. Interestingly,
we also found that Yapu women expressed a
preference for cultivars high in starch, but
not too high, as roots with a very high starch
content are physically hard and hence more
energy-demanding to grate, an important
step in detoxification.Third, Wilson ad-
dressed the question of differences between
high-CNP and low-CNP cultivars in suscept-
ibility to predation and disease in 1994 by
examining samples of plants (above- and
8 D.L. DUFOUR
below-ground portions) for evidence of
pathogen and pest damage. He did not find
significant differences in damage to the
above-ground portions of the plants but did
find that pest/pathogen damage to high-CNP
roots tended to be less, but not significantly
less, than that for low-CNP roots (Wilson,
1997). Furthermore, informants consis-
tently reported that a common rodent
(Dacyprocta fulignosa) preferred the roots
of low-CNP manioc and preyed on them
more often; we were not able to quantita-
tively evaluate that claim.
Fourth, in terms of the suitability of high-
CNP versus low-CNP for certain foods we
found that Tukanoans explicitly stated prefer-
ences for the taste and texture of their staple
foods (casabe, fariña, and manicuera) made
with high-CNP cultivars. All of these products
were routinely made with high-CNP but could
be made with either high- or low-CNP roots and
were occasionally made with the latter. The
time and effort required to make them was the
same, as were the storage qualities. We were
easily able to distinguish the manicuera (by
taste) and the fariña (by taste and texture)
made with high-CNP as opposed to low-CNP
roots but not the casabe. The Tukanoans,
however, were the connoisseurs and perceived
differences in the taste and texture of casabe
made from high-CNP as opposed to low-CNP
roots in a blind triangle taste test (Dufour,
1993). The taste, texture, and color (whiteness)
are features that distinguish casabe in the
region and are markers of identity.
In addition to testing the above explana-
tions, Wilson used pile-sort exercises to help
define Tukanoan preferences. These exercises
suggested that the foods to be made from
manioc roots were very important in the pre-
ference for high-CNP manioc cultivars and
were more important than either yield or
susceptibility to pest and pathogen damage
(Wilson and Dufour, 2002). These results
seem counterintuitive because our bias is to
think of yield as the pre-eminent factor in the
selection of food crops. Not all peoples,
however, place such emphasis on yield (Bos-
ter, 1984).
Taken together, these results suggest that
the Tukanoan preference for high-CNP man-
ioc is a complex interaction between local
ecological factors favoring cultivars with high-
CNP, human perceptions of and behavioral
responses to cyanogens in the staple crop, and
adequate biological responses to residual
cyanogens in processed foods. These interac-
tions are molded by an ideational system and
gustatory responses (learned?) that also favor
the more cyanogenic varieties of the staple
crop; a positive feedback loop. Hence, Tu-
kanoans are not merely responding behavio-
rally and biologically to the stress of
cyanogens in the environment, as they might
with a wild food resource; they are actually
manipulating the environment in ways that
increase the presence of cyanogens in the food
chain. The complexity of interactions revealed
by the studies described above demonstrate
the value of going beyond the usual and
measuring multiple causal pathways. This
complex set of interactions is no doubt the end
result of a long learning process by the
inhabitants of Amazonia.
CONCLUSIONS
Biocultural approaches have become an
integral part of research in human biology
and biological anthropology. They have,
however, tended to emphasize the ‘‘bio’’
and have not always risen to the challenge
of adequately addressing the ‘‘cultural.’’ I
hope that the foregoing examples have
illustrated the importance of focusing more
attention on the cultural phenomena critical
to our understanding of key features of
human biology. I also hope the examples
have illustrated the kind of dynamic inter-
actions among biological, cultural, and eco-
logical phenomena we should expect to find.
ACKNOWLEDGMENTS
I thank Barbara Piperata, Warren Wilson,
and Paul Patmore for their thoughtful
comments on this written version of the
Pearl Memorial Lecture, and Mike Little for
suggestions about topics to include. I also
thank friends in Cali, as well as Yapu, for
their goodwill and patience in teaching me a
bit about their world.
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