Pulmonary nitric oxide in mountain dwellers.
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Adaptation, Physiological↗
Biomedical subjects
C M Beall
Publications and source records attributed to C M Beall.
Tibetan and Andean contrasts in adaptation to high-altitude hypoxia.
High-altitude environments provide natural experimental settings to investigate adaptation to environmental stress. An important evolutionary and functional question is whether sea-level human biology constrains the adaptive response. This paper presents evidence that indigenous populations of the Tibetan and Andean plateaus exhibit quantitatively different responses to hypobaric hypoxic stress. At the same altitude, Tibetan mean resting ventilation and hypoxic ventilatory response were more than one-half standard deviation higher than Andean Aymara means while Tibetan mean oxygen saturation and hemoglobin concentration were more than one standard deviation below the Andean means. Quantitative genetic analyses of the familial patterning of these traits provided indirect evidence of population differences in genes influencing them. The Tibetan and Andean patterns of oxygen transport appear equally effective functionally as evaluated by birthweight and maximal aerobic capacity across a range of altitudes.
Acclimatization↗
Oxygen saturation increases during childhood and decreases during adulthood among high altitude native Tibetians residing at 3,800-4,200m.
This report describes age differences in oxygen saturation throughout the life cycle in a sample of high altitude native Tibetans residing in villages at 3,800-4,200 m altitude in the Tibet Autonomous Region, China. Oxygen saturation of 3,812 Tibetans was measured by pulse oximetry and a subsample of 1,582 healthy, nonpregnant, nonsmokers from 1 week to 80 years of age was selected for analyses. Infants under 1 year of age had 5-6% lower oxygen saturation than the peak of 89.8% attained at 11 years of age. There was a steady increase in mean oxygen saturation-for-age during the first decade of life, but not during the second decade. Adult males exhibited a slight decrease starting in the 20-29 year age range. Adult females maintained the peak oxygen saturation through the 40-49 year age range, exhibiting a decrease in oxygen saturation beginning in the 50-59 year age range and as a result had higher oxygen saturation than males during the female reproductive span. Thus, developmental factors during infancy and childhood, but not adolescence, enhanced oxygen transfer in this high altitude native resident Tibetan sample. The age of onset of aging processes detrimental to oxygen transfer differed for females and males.
Adolescent↗
Tibetan and Andean patterns of adaptation to high-altitude hypoxia.
Understanding the workings of the evolutionary process in contemporary humans requires linking the evolutionary history of traits with their current genetics and biology. Unusual environments provide natural experimental settings to investigate evolution and adaptation. The example of high-altitude hypoxia illustrates some of the progress and many of the remaining challenges for studies of evolution in contemporary populations. Current studies exemplify the frequently encountered problem of determining whether large, consistent population differences in mean values of a trait reflect genetic differences. In this review I describe 4 quantitative traits that provide evidence that indigenous populations of the Tibetan and Andean plateaus differ in their phenotypic adaptive responses to high-altitude hypoxia. These 4 traits are resting ventilation, hypoxic ventilatory response, oxygen saturation, and hemoglobin concentration. The Tibetan means of the first 2 traits were more than 0.5 standard deviation higher than the Aymara means, whereas the Tibetan means were more than 1 standard deviation lower than the Aymara means for the last 2 traits. Quantitative genetic analyses of within-population variance revealed significant genetic variance in all 4 traits in the Tibetan population but only in hypoxic ventilatory response and hemoglobin concentration in the Aymara population. A major gene for oxygen saturation was detected among the Tibetans. These findings are interpreted as indirect evidence of population genetic differences. It appears that the biological characteristics of sea-level humans did not constrain high-altitude colonists of the 2 plateaus to a single adaptive response. Instead, microevolutionary processes may have operated differently in the geographically separated Tibetan and Andean populations exposed to the same environmental stress. Knowledge of the genetic bases of these traits will be necessary to evaluate these inferences. Future research will likely be directed toward determining whether the population means reflect differences identified at the chromosomal level. Future research will also likely consider the biological pathways and environmental influences linking genotypes to phenotypes, the costs and benefits of the Tibetan and Andean patterns of adaptation, and the question of whether the observed phenotypes are indeed adaptations that enhance Darwinian fitness.
Adaptation, Physiological↗
Percent of oxygen saturation of arterial hemoglobin among Bolivian Aymara at 3,900-4,000 m.
A range of variation in percent of oxygen saturation of arterial hemoglobin (SaO2) among healthy individuals at a given high altitude indicates differences in physiological hypoxemia despite uniform ambient hypoxic stress. In populations native to the Tibetan plateau, a significant portion of the variance is attributable to additive genetic factors, and there is a major gene influencing SaO2. To determine whether there is genetic variance in other high-altitude populations, we designed a study to test the hypothesis that additive genetic factors contribute to phenotypic variation in SaO2 among Aymara natives of the Andean plateau, a population geographically distant from the Tibetan plateau and with a long, separate history of high-altitude residence. The average SaO2 of 381 Aymara at 3,900-4,000 m was 92+/-0.15% (SEM) with a range of 84-99%. The average was 2.6% higher than the average SaO2 of a sample of Tibetans at 3,800-4,065 m measured with the same techniques. Quantitative genetic analyses of the Aymara sample detected no significant variance attributable to genetic factors. The presence of genetic variance in SaO2 in the Tibetan sample and its absence in the Aymara sample indicate there is potential for natural selection on this trait in the Tibetan but not the Aymara population.
Adaptation, Physiological↗
Hemoglobin concentration of high-altitude Tibetans and Bolivian Aymara.
Elevated hemoglobin concentrations have been reported for high-altitude sojourners and Andean high-altitude natives since early in the 20th century. Thus, reports that have appeared since the 1970s describing relatively low hemoglobin concentration among Tibetan high-altitude natives were unexpected. These suggested a hypothesis of population differences in hematological response to high-altitude hypoxia. A case of quantitatively different responses to one environmental stress would offer an opportunity to study the broad evolutionary question of the origin of adaptations. However, many factors may confound population comparisons. The present study was designed to test the null hypothesis of no difference in mean hemoglobin concentration of Tibetan and Aymara native residents at 3,800-4,065 meters by using healthy samples that were screened for iron deficiency, abnormal hemoglobins, and thalassemias, recruited and assessed using the same techniques. The hypothesis was rejected, because Tibetan males had a significantly lower mean hemoglobin concentration of 15.6 gm/dl compared with 19.2 gm/dl for Aymara males, and Tibetan females had a mean hemoglobin concentration of 14.2 gm/dl compared with 17.8 gm/dl for Aymara females. The Tibetan hemoglobin distribution closely resembled that from a comparable, sea-level sample from the United States, whereas the Aymara distribution was shifted toward 3-4 gm/dl higher values. Genetic factors accounted for a very high proportion of the phenotypic variance in hemoglobin concentration in both samples (0.86 in the Tibetan sample and 0.87 in the Aymara sample). The presence of significant genetic variance means that there is the potential for natural selection and genetic adaptation of hemoglobin concentration in Tibetan and Aymara high-altitude populations.
Adaptation, Physiological↗
Ventilation and hypoxic ventilatory response of Tibetan and Aymara high altitude natives.
Newcomers acclimatizing to high altitude and adult male Tibetan high altitude natives have increased ventilation relative to sea level natives at sea level. However, Andean and Rocky Mountain high altitude natives have an intermediate level of ventilation lower than that of newcomers and Tibetan high altitude natives although generally higher than that of sea level natives at sea level. Because the reason for the relative hypoventilation of some high altitude native populations was unknown, a study was designed to describe ventilation from adolescence through old age in samples of Tibetan and Andean high altitude natives and to estimate the relative genetic and environmental influences. This paper compares resting ventilation and hypoxic ventilatory response (HVR) of 320 Tibetans 9-82 years of age and 542 Bolivian Aymara 13-94 years of age, native residents at 3,800-4,065 m. Tibetan resting ventilation was roughly 1.5 times higher and Tibetan HVR was roughly double that of Aymara. Greater duration of hypoxia (older age) was not an important source of variation in resting ventilation or HVR in either sample. That is, contrary to previous studies, neither sample acquired hypoventilation in the age ranges under study. Within populations, greater severity of hypoxia (lower percent of oxygen saturation of arterial hemoglobin) was associated with slightly higher resting ventilation among Tibetans and lower resting ventilation and HVR among Aymara women, although the associations accounted for just 2-7% of the variation. Between populations, the Tibetan sample was more hypoxic and had higher resting ventilation and HVR. Other systematic environmental contrasts did not appear to elevate Tibetan or depress Aymara ventilation. There was more intrapopulation genetic variation in these traits in the Tibetan than the Aymara sample. Thirty-five percent of the Tibetan, but none of the Aymara, resting ventilation variance was due to genetic differences among individuals. Thirty-one percent of the Tibetan HVR, but just 21% of the Aymara, HVR variance was due to genetic differences among individuals. Thus there is greater potential for evolutionary change in these traits in the Tibetans. Presently, there are two different ventilation phenotypes among high altitude natives as compared with sea level populations at sea level: lifelong sustained high resting ventilation and a moderate HVR among Tibetans in contrast with a slightly elevated resting ventilation and a low HVR among Aymara.
Acclimatization↗
Quantitative genetic analysis of arterial oxygen saturation in Tibetan highlanders.
This study was designed to test the hypothesis that genetic differences inferred from biological kinship relationships among individuals contribute to individual variation in percentage of oxygen saturation of arterial hemoglobin (SaO2) in a high-altitude native population. SaO2 data were obtained by pulse oximetry from 354 nonpregnant, healthy Tibetan residents of Pen-Dri, two rural agropastoral villages at 3800-4065 m altitude in Lhasa Municipal District, Tibet Autonomous Region, China. Statistical analyses of these data from 46 pedigrees tested the hypothesis of a significant genetic contribution to SaO2 variation. The average SaO2 was 89.4 +/- 0.2%, with a range of 76-97%. Additive genetic effects account for 44% of the interindividual phenotypic variation in SaO2 in the sample. Complex segregation analysis and variance decomposition analysis determined that 21% of the total phenotypic variation could be explained by a major gene influencing SaO2. Homozygotes for the low-SaO2 allele have a mean SaO2 of 83.6%, whereas heterozygotes and homozygotes for the high-SaO2 allele have means of 87.6% and 88.3%, respectively. This confirms findings in another Tibetan sample and extends the known geographic distribution of the major gene. These results suggest the hypothesis that individuals with the dominant allele for higher SaO2 have a selective advantage in their high-altitude hypoxic environment.
Adaptation, Physiological↗
Major gene for percent of oxygen saturation of arterial hemoglobin in Tibetan highlanders.
This report employs a statistical genetic approach to analyze quantitative oxygen transport variables in a high-altitude (4,850-5,450 m) native Tibetan population and demonstrates the presence of a major gene influencing % O2 saturation of arterial hemoglobin. This result suggests the hypothesis that individuals with the dominant allele for higher % O2 saturation have a selective advantage at high altitude. Studies of the biologically distinctive Himalayan and Andean populations have greatly influenced thinking about ongoing human evolution and adaptation; this is the first statistical evidence for a major gene enhancing oxygen transport in a high-altitude native population.
Adaptation, Physiological↗
Hemoglobin concentration of pastoral nomads permanently resident at 4,850-5,450 meters in Tibet.
This paper presents data on the hemoglobin concentration of a sample of 103 pastoral nomads who are lifelong residents of Phala, at 4,850-5,450 m, on the northern plateau of the Tibet Autonomous Region of the Peoples' Republic of China. This native population resides at the highest altitude of which we are aware and is thus exposed to the most extreme chronic hypoxic stress. However, they do not exhibit the most pronounced physiological adaptations, i.e., hemoglobin concentrations exceeding those found in all other high-altitude populations. Adult male and female mean hemoglobin concentrations of 18.2 and 16.7 gm/dl, respectively, were found. These data, in conjunction with earlier studies of ethnic Tibetans living at 3,400 m, demonstrate a pattern of increasing hemoglobin concentration (erythrocytosis) at increasing altitude of residence in the Himalayas and Tibet. At the same time, however, the hemoglobin concentration is lower than that found among Andean highlanders. These new data raise the possibility of quantitative population differences in hematological adaptation to high altitude hypoxia.
Adolescent↗
Age differences in sensory and cognitive function in elderly Nepalese.
This paper describes age differences in sensory and cognitive function in a cross-sectional sample of 117 Nepalese men aged 50 to 88 years living in a traditional agrarian society. The prevalence of impairment of vision, hearing, vibration sensitivity, and cognitive function is progressively higher in successively older age categories in this society as it is in Western industrial societies. This has practical consequences for everyday life. Men with vision impairment are less likely to retain the esteemed social role of head of an extended household. Men with visual and hearing impairment leave their household compounds less frequently, have less frequent social contacts outside their households, and remain inactive a greater proportion of the time. In this technologically simple society without modern medical remedies, vision and hearing impairments are associated with social roles and daily activity patterns that foster economic and political dependence and social isolation.
Aged↗
The physical fitness of elderly Nepalese farmers residing in rugged mountain and flat terrain.
This paper reports the results of a natural experimental test of the hypothesis that elderly residents of rugged terrains are more physically fit than their counterparts in flat terrains due to the additional lifelong daily exertion occasioned by the inescapable need to walk up and down steep slopes. Cycle ergometer tests of physiological response to work were undertaken by two groups of high caste Hindu male farmers, aged 50 to 79, living in Central Nepal. One comprised an experimental group resident in rugged terrain and the other a control group resident in flat terrain. No differences in heart rate and blood pressure measured at submaximal workloads, at peak effort, or during recovery from exercise were demonstrated. Measurements of physical exertion revealed that few people in either terrain engaged in activity sufficiently strenuous to raise heart rates to a level where training could occur. Despite a rural agricultural lifestyle, these men were not especially physically fit.
Aged↗
Social structure and intracohort variation in physical fitness among elderly males in a traditional Third World society.
This paper examines the relationship between physical fitness and activity among elderly males in the traditional rural community of Chetbesi, Nepal. It takes advantage of the unique character of the Hindu caste system to implement a quasiexperimental research design that approximates random assignment to high and low activity levels. The members of the Sarki caste have lower heart rates and systolic blood pressure, relative to other castes, at each of three submaximal workloads and during recovery from bicycle ergometer exercise. Direct observation and physiologic monitoring show that the Sarkis engage in more frequent and extended periods of heavy labor. Thus intracohort variation in physical fitness and activity patterns among the Chetbesi elderly is a function of birth into a socially defined group rather than of self-selection. This pattern of differential fitness may typify the process of aging in many stratified traditional and modernizing societies where socially delimited segments of the population perform the bulk of the hard work. Intrapopulation differences aside, comparison of Sarkis and non-Sarkis with other samples reveals that both lie within the reported range of variation. The rural, unmechanized, agricultural lifestyle and mountain environment of Chetbesi do not result in exceptional fitness for residents. The Chetbesi data suggest that the popular notion that aging is less debilitating in traditional agrarian societies located in rugged mountain terrains may be a myth. The demonstration of the influence of social forces on physical fitness suggests that future research might concentrate profitably on identifying social structures that produce high levels of physical fitness.
Aged↗
Hemoglobin levels in a Himalayan high altitude population.
This report presents data on hemoglobin concentrations in a sample of Himalayan high altitude natives measured at their habitual altitude of residence. In this sample of 270 healthy Tibetan adults resident at 3250-3560 m in Upper Chumik , Nepal, the mean hemoglobin concentration is 16.1 +/- 1.2 gm/dl among adult males, 14.4 +/- 1.4 gm/dl among premenopausal and 15.0 +/- 1.1 gm/dl among postmenopausal adult females. 123 of 126 (98%) males, 96 of 100 (96%) premenopausal and 36 of 44 (82%) postmenopausal females have hemoglobin concentrations within two standard deviations of the sea level mean. These data demonstrate that a healthy population may reside at high altitude without the degree of elevation in hemoglobin widely known and cited for Andean highlanders. Comparing published data on mean hemoglobin concentrations of adult Himalayan and Andean samples residing between 3200 m and 4100 m reveals that Himalayan means are systematically lower. This in turn may account for the reported population differences in the prevalence of chronic mountain sickness ( Monge 's disease). It is hypothesized that Himalayan and Andean highlanders represent alternative patterns of high altitude hematological adaptation.
Adaptation, Physiological↗
Cross-cultural studies of biological aging. Introduction.
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Aging↗
Biological function, activity and dependency among elderly Sherpa in the Nepal Himalayas.
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Aging↗
Work, aging and dependency in a Sherpa population in Nepal.
This paper suggests a conceptual framework for the cross-cultural study of dependency in order to encourage and facilitate data-based cross-cultural comparisons. It offers an etic, operational definition encompassing biological, activity and economic components. It also emphasizes that the emic definition is crucial for understanding a situation. The utility of distinguishing several components of an etic definition of dependency and distinguishing etic from emic definition is illustrated using the examples of Sherpas in Helambu, Nepal. From the etic viewpoint, the Sherpas are apparently healthy, economically productive and physically active and therefore lie near the independent end of the dependency continuum. The recent emigration of many young adults has produced a demographic situation where few elderly can achieve their expectation of growing old while sharing their own households with their youngest son and his family. Although family life is valued and most elderly Sherpas have other offspring and other relatives in the village, the majority of elderly Sherpa live alone and express dissatisfaction at doing so. From their standpoint, reliance upon someone other than the youngest son for lodging or food is considered prejoratively and as dependency. Their emic definition of dependency is different from the Western definition and both differ from what we have referred to as etic dependency.
Aged↗
Indirect modernization and the status of the elderly in a rural Third World setting.
Currently the majority of the world's elderly reside in less developed countries and their proportion is increasing. This paper presents evidence that the process of modernization can have a negative impact on the elderly in even the most remote rural Third World settings, even though those settings are not modernized or in the process of modernization in any of the normal uses of that concept. Fieldwork was conducted in Helambu, Nepal on a sample of 37 persons over the age of 50 that included 86% of population aged 60 and over. Despite high levels of activity, health, social and economic status, the elderly were greatly dissatisfied with their situation. The paper demonstrates the manner in which modernization in India has profoundly changed household/family organization in Helambu and produced this situation.
Aged↗