AMERICAN JOURNAL OF HUMAN BIOLOGY 18:19 (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 interactions 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 explanations 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 ethnographically 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 # 2005 Wiley-Liss, Inc. energetics as well as that of other practitioners. Am. J. Hum. Biol. 18:19, 2006.

Human biology is an interdisciplinary field centrally concerned with understanding human biological variability and the mechanisms responsible for that variability. Researchers represent a variety of perspectives, particularly those that are comparative, developmental, ecological, and/or evolutionary. Within each of these perspectives, biocultural approaches are those that explicitly recognize the dynamic interactions between humans as biological beings and the social, cultural, and physical environments they inhabit. They focus on understanding variability in human biology as a function of responsiveness to the larger (social, cultural, and physical) environment. Biocultural approaches have a long history in human biology and biological anthropology, a closely linked scientific community. They can, however, be very challenging approaches to implement, perhaps in part because we are more experienced in measuring the biological than the social or cultural. Some of the challenges include (1) defining precisely what we mean by constructs like socioeconomic status, poverty, social support, etc.; (2) operationalizing key variables so that they can be measured in ways that are replic-

able as well as ethnographically valid; and (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.

A BRIEF HISTORY It is difficult to know where to begin, but because this is the Pearl Memorial Lecture, it is fitting that we start with Raymond Pearl. Although Pearl is best known for his contributions to biostatistics, he clearly understood the impact of the social and cultural environment on human biology and employed culturally defined variables in many of his analyses. The importance Pearl attributed to the social and cultural environment is clear in the following passage:
*Correspondence to: Darna L. Dufour. E-mail: Darna. Dufour@colorado.edu Received 29 August 2005; Accepted 19 September 2005 Published online in Wiley InterScience (www.interscience. wiley.com). DOI: 10.1002/ajhb.20463

2005 Wiley-Liss, Inc.

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 environment on human biology, at least in comparison 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 evolution 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 conceptualize 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 linkages 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. Livingstones contribution notwithstanding, the dominant idea remained, however, to consider the environment as the physical environment. This is clear in the first textbook in human biology, the 1964 text Human BiologyAn Introduction to Human Evolution, 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 communications; the vegetational cover and the insect animal and bird life . . . The natural environment 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 biologically 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 limited to the physical environment characterized the early stages of the human adaptability paradigm. If we assume, with Kuhn (1970), that textbooks 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 cultural 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 different 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 demonstrated the potential of a more holistic view of human populations. It probably served as the template for Morans (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 environment. It did, however, include a chapter on the westernization of dietary habits and disease expression that has a biocultural focus. It was around the same time that the term biocultural started to appear in the literature. The first paper in the journal Human

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Biology with the term biocultural in the title was Biocultural correlates of blood pressure 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 chapters 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 modernization, a culturally defined construct, on biology (Baker 1986). The Samoan research exemplified the deepening interest among biological anthropologists in culturally defined independent variables and a willingness to consider the social and cultural components of the environment.The most recent textbook, Human Biology: An Evolutionary and Biocultural Perspective, edited by Stinson et al. (2000) was written collaboratively by members of the Human Biology Association. The book includes the term biocultural in the title and in the Preface; the editors state:Human biology relies heavily on an evolutionary perspective to explain variation through space and time but also considers to be crucial the effect that human cultures have on our biologya biocultural perspective. (Stinson et al., 2000)The chapters 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 biocultural 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 Leatherman (1998). In particular, the editors place a

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 understand 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 important elements in determining the biologically significant environment of human beings . . . (Pearl, 1930:540). CHALLENGES Biocultural approaches can be very challenging 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 variables 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 wellbeing, 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,

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social exclusion, humiliation, and dependency (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 developing 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 specifically 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 peoples 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 midlow, and a high) following the definitions 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 undernutrition during growth and development. SES differences in BMI, were not, however, what we expected to find. First, there were very few women (35%) 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 highSES group. Third, twice as many of the lowSES 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 overweight 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 lowSES group was a question begging for a biocultural approachone with real attention to the cultural component.The approach I took was an inductive one: to try to understand the links between poverty and nutritional status in this particular ethnographic context. As a start, I spent some time doing ethnographic observations and informal interviews. Those initial ethnographic 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 storagenot even of things like salt and sugar; food was purchased on a mealto-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 minimal, 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: cows 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-

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tion because the foods were socially inappropriate. As the study progressed over the next 4 years or so, these initial impressions were confirmed by further observations and interviews. 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 requirement; 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 26 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 dieting (a very rare behavior in this population). Hence, we assumed that these lowintake days represented economic constraint 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. Shortterm here is day-to-day for most households, week-to-week for others. Some of the fluctuations 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 dont have money you dont 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 26 low-intake days had significantly lower energy intakes (2273 versus 1833 kcal/day) and activity levels (total daily energy expenditure (TDEE)/BMR 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,

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 (18.525 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 dimension 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, demographic, epidemiological, nutritional, etc., as well as characteristics of the human-built and physical environments) should lead to differences of interest. Hence, we should be thinking 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 biological 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 independent 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

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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 ethnographic understanding of the local setting. This is a point Dressler (1995) has emphasized. The survey-type interview data we collect does not really go deep enough to give us the kind understanding of the local ethnographic 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 energetics of working versus not-working women 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 understanding 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 income, no matter the time commitment per day, or whether it was short or long term, occasional, steady, or even highly unpredictable. For example, some women felt that they worked for the research project because they were remunerated for their participation 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 example, 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. Geertzs (1973) notion of thick description applies here; we need a thick understanding. Without it, we end up doing a remote-sensing kind of fieldwork without 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 interested in understanding complex interactions between biology and culture, then the consideration of multiple variables is invaluable. 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 preference for manioc (cassava; Manihot esculenta 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 potential (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,

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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 cultivars were toxic and correctly associated the toxicity with a bitter taste and an acrid smell (Dufour, 1988). Nonetheless, these highCNP 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 detoxification (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 techniques) and culturally (considering the foodways 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 cultivated primarily the high-CNP varieties, which they referred to as kii. Tukanoan women each explained it in terms of tradition: their mother did it that way, and their mothers 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

introduced by rubber gatherers from Brazil around the 1940s. These explanations were intuitively appealing because the importance of history and tradition in a group such as this cannot be denied. There was, however, one intriguing incongruity: lowCNP manioc was introduced at about the same time as a group of yellow-fleshed highCNP 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 a (manioc meal), which used to make farin was introduced at the same time. So tradition 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 current 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 glucosides 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 literature 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 significant in the 1994 data (Dufour, 1993; Wilson and Dufour, 1996, 2002).Second, in terms of starch content, our results were contradictory: 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 addressed the question of differences between high-CNP and low-CNP cultivars in susceptibility to predation and disease in 1994 by examining samples of plants (above- and

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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 consistently 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 quantitatively evaluate that claim. Fourth, in terms of the suitability of highCNP versus low-CNP for certain foods we found that Tukanoans explicitly stated preferences for the taste and texture of their staple a, and manicuera) made foods (casabe, farin 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 a (by taste and texture) taste) and the farin 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 explanations, 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 preference 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 (Boster, 1984). Taken together, these results suggest that the Tukanoan preference for high-CNP manioc is a complex interaction between local ecological factors favoring cultivars with highCNP, human perceptions of and behavioral responses to cyanogens in the staple crop, and adequate biological responses to residual

cyanogens in processed foods. These interactions 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, Tukanoans are not merely responding behaviorally 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 interactions among biological, cultural, and ecological 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. LITERATURE CITED
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