The home team advantage: reproduction in women indigenous to high altitude.

Although there is substantial evidence that environmental conditions disrupt reproductive function among newcomers to hypoxic settings, it is not certain that low oxygen pressure reduces fertility among those indigenous to high altitude. Even when fertility does appear to be relatively lower, numerous behavioral and sociocultural factors may be responsible. These are best examined within demographic frameworks that delineate a finite list of the proximate determinants of fertility. The findings presented here are based on several studies of indigenous Andean populations (Peruvian Quechua at 4000 m, Bolivian Quechua at 3100 m, Bolivian Aymara at 4000 m). Data on ovarian function suggest that neither progesterone levels nor menstrual cycle length or regularity are significantly different from those of women at lower altitudes. Data on two behavioral factors that determine fertility levels, coital frequency and infant feeding practices, suggest that the former is not likely to be of significance in co-habitating couples, but that variation in breastfeeding patterns has probably made a substantial contribution to differences in fertility among at least some populations at high altitude.

Vaginal bleeding patterns among rural highland Bolivian women: relationship to fecundity and fetal loss.

Among the most common reasons given for discontinued use of some contraceptive methods is a disturbance in the menstrual cycle, particularly changes in vaginal bleeding. Work to date suggests marked populational variation in menses duration, but few data have been collected from South America. This longitudinal study of non-contracepting Aymara women (n = 189 providing 837 non-truncated bleeding episodes) identified conceptions and fetal loss via urine tests for human chorionic gonadotropin and classified episodes accordingly to test the hypotheses that (a) vaginal bleeding patterns differ between lactating and non-lactating women, (b) duration of vaginal bleeding accompanying fetal loss differs from that of menstruation, (c) menses preceding a conception are longer than those not followed by a conception. Compared to published values, mean menses duration (3.5 days) in these women was relatively short. Menses duration was not significantly correlated with current age, age at menarche or first birth, parity, time postpartum, or menstrual segment length. Mean menses duration (not preceding a conception) was comparable for lactating and non-lactating women. Mean duration of fetal loss bleeding did not differ from that of menses. Pre-conception episodes were significantly longer than those not followed by conception. Thus, because the rate of conceptions was twice as great among lactating than non-lactating women, the mean duration of all menses (irrespective of conception) was significantly longer in lactating women. Bolivian, and perhaps other South American, women may be particularly disinclined to accept contraceptives (e.g., intrauterine devices) that modify an otherwise relatively brief menses duration. Therefore, a wide variety of contraceptive choices accompanied by population-specific informed counseling is essential. In addition, these findings suggest that studies of fecundability limited to non-lactating women may be biased toward those of relatively lower fecundity and that menses duration may be predictive of risk for some cancers.

Menstrual patterns and fecundity among non-lactating and lactating cycling women in rural highland Bolivia: implications for contraceptive choice.

Choosing an appropriate contraceptive method, particularly one based on fertility awareness, depends in part upon the degree of a woman's cycling regularity. However, while the suppressive effect of lactation on ovarian function is well established, the potential influence of continued breastfeeding on menstrual patterns once post-partum cycling has resumed is largely unexamined. This longitudinal study in a population of non-contracepting Aymara women (n = 191 providing 665 non-truncated menstrual segments) identified conceptions and fetal loss via urine tests for hCG and classified segments accordingly to test the hypotheses that (1) cycles in lactating women are significantly different in length and regularity from those of non-lactating women, and (2) cycles in women living at high altitude are significantly different in length and regularity from those at lower altitudes. Analyses found that segments are significantly longer and regularity tends to be less common among lactating than non-lactating women; however, the rate of conception is twice as great among the former than the latter, and the distribution of conception outcomes does not differ between the two groups. Menstrual regularity is not the norm in this population, even among those who are not currently breastfeeding. High altitude per se does not appear to influence menstrual cycling as both regularity and segment length are comparable to other populations, hence women indigenous to high altitude are suitable candidates for a wide variety of contraceptive choices. In addition, these findings suggest that studies of fecundability limited to non-lactating women may be biased towards those of relatively lower fecundity.

Does hypoxia impair ovarian function in Bolivian women indigenous to high altitude?

Fertility appears to be reduced in at least some high altitude populations relative to their counterparts at lower elevations. Inferring from the difficulties with reproduction of newcomers to high altitude and from animal experiments, it has been hypothesized that this apparent reduction is the result of hypoxia acting to reduce fecundity and/or increase fetal loss. In humans, however, several behavioral as well as biological factors may affect fertility levels. These many factors have been organized by demographers into a framework of seven proximate determinants that includes fecundability (the monthly probability of conception) of which successful ovulation is one component. To test whether ovarian function is impaired in women indigenous to high altitude, we measured salivary progesterone (P) in a sample (n = 20) of Quechua women (aged 19-42 years) residing at 3,100 m. It was found that mean luteal P = 179 pmol/L and mean midluteal P = 243 pmol/L, levels that fall about midway in the range of known values for several populations and are higher than some lower altitude populations. These findings suggest that hypoxia does not appear to significantly impair ovarian function in those with lifelong residence at high altitude. There are, however, several factors common to many high altitude populations that may act to reduce fecundability and fertility including intercourse patterns (affected by marriage and migration practices), prolonged lactation, dietary insufficiency, and hard labor.

Causes and consequences of heterogeneity in infant feeding practices among indigenous Andean women.

PIP: Population variation in maternal work schedules milk output, presence of surrogate caretakers, supplemental feeding patterns, and maternal-infant co-sleeping very likely affects suckling duration and frequency. There can be large differences, even in relatively homogenous rural indigenous populations that affect fertility, maternal-infant health, and intervention programs. Findings were presented on suckling patterns and on heterogeneity in fecundity and socioeconomic status (SES) and feeding practices among three rural Quechua communities in Nunoa, Peru. Mothers and infants were from low SES and mid SES households. The suckling time to the nearest second among 10 infants showed longer and more frequent nursing sessions in the morning and reduced feeding during the night. Duration of suckling was associated with infant's age, and declined with increasing age of the infant. Interval duration was not associated with infant's age. Mother's age was found to be associated with afternoon suckling duration. Younger mothers had shorter afternoon sessions, perhaps because there were older adolescent women who cared for older mothers' children in the afternoon or older mothers produced less milk and thus required longer suckling times. Afternoon interval duration did not vary with infant's or mother's age. Maternal recall of suckling duration and frequency was considered inaccurate, as a result of recall and observational comparisons of duration and frequency for a sample of 9 women. Postpartum amenorrhea averaged 18.8 months for median duration. Mid SES women had a postpartum amenorrhea period of 8.8 months compared to 21.6 months for low SES women. Mid SES women had shorter breast feeding durations during the day of 16.6 months compared to 23.6 months for low SES women, and during the night, 2.4 years versus 3.6 years. About 40% of both low and mid SES women used bottle feeding and began bottle feeding at the same age of 5 months. Solid food was introduced at 13 months for both groups of women.^ieng

Brief communication: effect of coca-leaf chewing on salivary progesterone assays.

Although there is evidence for reduced fertility in Andean and Himalayan populations at higher altitudes, factors other than hypoxia may be primarily responsible. A valuable approach in the investigation of these fertility determinants is the use of salivary steroid assays. However, coca-leaf chewing--a ubiquitous practice among high altitude Andean populations--has negative consequences for the accurate measurement of ovarian steroids. This report evaluates the effects of coca-leaf chewing on assays of salivary progesterone. Study participants include naive and habitual users of coca leaf from La Paz and El Alto, Bolivia. Approximately 300 saliva samples were collected immediately before, during, and after coca-leaf chewing. The series includes samples with and without the alkaloid enhancer typically used by coca-leaf chewers. Coca chewing produces false salivary progesterone values that mimic luteal phase values. On the basis of this study, an appropriate protocol is developed for the collection of salivary samples in coca-leaf chewing populations. These results verify the feasibility of salivary assays, even for very difficult field conditions, and highlight the necessity of establishing suitable collection procedures before full field implementation of saliva sampling.

Infant nutrition and the consequences of differential market access in Nunoa, Peru.

PIP: The author observed and interviewed 30 women in August 1985, each of whom had at least 1 child under 3 years old, to analyze infant feeding behavior in the rural Andean community of Nunoa, Peru. Infants were generally first nursed within 48 hours of birth, subject to co-sleeping and probably night-nursing far longer than day nursing, and fed exclusively via bottle or nursing on only very rare occasions. Weaning periods were extended and consisted of continually modified feeding routines. Significant differences were observed between the feeding habits of poor and relatively less-poor mothers which may have impacted differently upon the health of offspring. The quality of infant foods and economic variation influenced breastfeeding duration and subsequent infant health and maternal fecundity. The author concludes that infant feeding routines can be neither characterized nor studied as a set of mutually exclusive hierarchical stages; significant variation exists within and between socioeconomic subsamples; both groups have parallel perceptions and attitudes regarding appropriate feeding practices; both groups perceive foods other than breast milk as complements to rather than substitutes for nursing during the 1st year of life; bottle use is neither uniformly harmful nor beneficial; the various aspects of infant feeding routines do not all necessarily respond in concert to changing socioeconomic conditions; models of infant feeding behavior must go beyond the simple mechanics of infant feeding; and great need exists for more comprehensive investigations.^ieng

Nursing behaviour and its relation to duration of post-partum amenorrhoea in an Andean community.

Nuñoa is a high altitude rural Peruvian community characterized by socioeconomic stratification and differential access to the market economy. Nursing practices and the effects of nursing are also stratified; this translates into a risk of menses at 12 months post-partum nearly seven times greater in wealthier than in poorer women. Most nursing occurs in the morning, among those who practice on-demand breast-feeding. Nursing episodes are clustered into sessions; the amount of breast-feeding is regulated by varying the number of episodes per session rather than by changing the duration of suckling episodes or the frequency of sessions per hour. Thus, resumption of ovulation is not dependent on the variable spacing of nursing episodes or sessions. The components of nursing activity most likely to be responsible for variation in the duration of post-partum amenorrhoea in these nursing women are mean session duration and mean number of episodes per session. The mean duration of morning nursing sessions is negatively associated with infant's age, reflecting the greater reliance of younger children on breast-milk. The mean duration of afternoon nursing sessions is positively associated with mother's age, independent of infant's age, possibly reflecting maternal age-related variation in milk production capabilities. Baby minding by older daughters may also help to explain variation in afternoon nursing.

Dental metric assessment of the omo fossils: implications for the phylogenetic position of Australopithecus africanus.

The discovery of Australopithecus afarensis has led to new interpretations of hominid phylogeny, some of which reject A. africanus as an ancestor of Homo. Analysis of buccolingual tooth crown dimensions in australopithecines and Homo species by Johanson and White (Science 202:321-330, 1979) revealed that the South African gracile australopithecines are intermediate in size between Laetoli/hadar hominids and South African robust hominids. Homo, on the other hand, displays dimensions similar to those of A. afarensis and smaller than those of other australopithecines. These authors conclude, therefore, that A. africanus is derived in the direction of A. robustus and is not an ancestor of the Homo clade. However, there is a considerable time gap (ca. 800,000 years) between the Laetoli/Hadar specimens and the earliest Homo specimens; "gracile" hominids from Omo fit into this chronological gap and are from the same geographic area. Because the early specimens at Omo have been designated A. afarensis and the later specimens classified as Homo habilis, Omo offers a unique opportunity to test hypotheses concerning hominid evolution, especially regarding the phylogenetic status of A. africanus. Comparisons of mean cheek teeth breadths disclosed the significant (P less than or equal to 0.05) differences between the Omo sample and the Laetoli/Hadar fossils (P4, M2, and M3), the Homo fossils (P3, P4, M1, M2, and M1), and A. africanus (M3). Of the several possible interpretations of these data, it appears that the high degree of similarity between the Omo sample and the South African gracile australopithecine material warrants considering the two as geographical variants of A. africanus. The geographic, chronologic, and metric attributes of the Omo sample argue for its lineal affinity with A. afarensis and Homo. In conclusion, a consideration of hominid postcanine dental metrics provides no basis for removing A. africanus from the ancestry of the Homo lineage.

The impact of random and lineal fission of the genetic divergence of small human groups: a case study among the Yanomama.

Most of the genetic divergence that currently separates populations of Homo sapiens must have arisen during that long period when the local village (or band) was the basic unit of biological evolution. Studies of tribally intact Amerindian groups exhibiting such small-group organization have demonstrated marked genetic divergence between nearby villages. Some of this genetic radiation can be attributed to the effects of random genetic drift over time within these small demes. Some of it, however, might be better ascribed to the consequences of nonrandom genetic assortment at the time of village fission, a recurring event for such groups. Even random genetic assortment at the time of fission would lead to some genetic divergence, due to the finite size of the parent gene pool. We term the genetic consequences of random assortment the random fission effect. Routinely, village fission occurs along family lines, leading to even greater genetic divergence between the daughter villages. We use the term lineal fission effect to describe the genetic consequences of nonrandom assortment and contrast these results with those derived from random assortment--A formal treatment of random and lineal fission effects is developed, first for the single-locus case, then for the multiple-locus extension. Using this formulation, three Yanomama fission events were examined. Fission in the Yanomama often involves a great deal of mutual hostility between the two factions, so that subsequent gene flow between the two daughter villages is minimal. The first two examples are typical of the Yanomama behavior norm, and are accompanied by a minimum of subsequent gene flow between the daughter villages. In these two cases, the observed divergence values are very large and are also very unlikely under random fission. The lineal fission effect is pronounced. The net impact of lineal fission is to reduce the effective size of the village at the time of fission by a factor of four, relative to expectation from random fission. The third example, however, involved an unusually amicable split of a village, followed by free genetic exchange between the fission products. This "friendly fission" yields an observed divergence value not much in excess of the expectation from random fission.--The long-term consequences of such fission bottlenecks in effective population size are discussed for both intra- and inter-tribal genetic diversity. It appears that the rate of genetic divergence for tribal and subtribal groups may have been somewhat greater than would be expected from classical drift arguments.