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. Am J Phys Anthropol 104:427–447, 1997. © 1997 Wiley-Liss, Inc.     

Responses of the autonomic nervous system in altitude adapted and high altitude pulmonary oedema subjects

Studies were carried out to ascertain the role of sympatho-parasympathetic responses in the process of adaptation to altitude. The assessment of status of autonomic balance was carried out in a group of 20 young male subjects by recording their resting heart rate, blood pressure, oral temperature, mean skin temperature, extremity temperatures, pupillary diameter, cold pressor response, oxygen consumption, cardioacceleration during orthostasis and urinary excretion of catecholamines; in a thermoneutral laboratory. The same parameters were repeated on day 3 and at weekly intervals for a period of 3 weeks, after exposing them to 3,500 m; and also after return to sea level. At altitude, similar studies were carried out in a group of 10 acclimatized lowlanders, 10 high altitude natives and 6 patients who had recently recovered from high altitude pulmonary oedema. In another phase, similar studies were done in two groups of subjects, one representing 15 subjects who had stayed at altitude (3,500–4,000 m) without any ill effects and the other comprising of 10 subjects who had either suffered from high altitude pulmonary oedema (HAPO) or acute mountain sickness (AMS). The results revealed sympathetic overactivity on acute induction to altitude which showed gradual recovery on prolonged stay, the high altitude natives had preponderance to parasympathetic system. Sympathetic preponderance may not be an essential etiological factor for the causation of maladaptation syndromes.     

Effect of altitude exposure on thermoregulatory response of man to cold.

The thermoregulatory responses to 10 degrees C (for 3 h) were investigated in 1) 12 natives from sea level (lowlanders) at 150 m, and on arrival at 3,350 and 4,340 m; 2) 6 of these during a 6-wk sojourn at 4,360 m, and on return to sea level; and 3) 5 natives from each of the two altitudes (highlanders) in their respective habitat, and after descent to 150 m. The cold-induced increase in the rate of O2 consumption (Vo2) of the lowlanders was significantly smaller at both altitudes than at sea level. It did not recover substantially during the 6 wk at altitude, but was restored to its initial rate on return to sea level. By contrast, visible shivering activity was augmented on arrival at altitude. It persisted throughout the 6 wk there, but was greatly depressed on return to sea level, despite the increased Vo2. Mean skin temperatures (Tsk) stabilized in the cold at significantly higher values at altitude. Rectal temperature (Tre) decreased similarly at all altitudes. Vo2 of the highlanders in the cold was significantly greater at sea level than at their resident altitudes, although shivering activity was less intense; Tsk stabilized at significantly lower levels at 150 m than at either altitude. These results indicate that altitude exposure reduces the calorigenic response of man to cold, and that this effect is not moderated by acclimatization to altitude, yet is reversible immediately on descent to sea level. The component of cold thermogenesis which appeared to be reduced by altitude exposure was nonshivering thermogenesis rather than visible shivering.     

Cell mediated immunity at high altitude

Cell mediated immunity (CMI) was assessed by determining total and differential leucocyte and absolute lymphocyte counts, T and B-rosettes, PHA-blast transformation of lymphocytes, lymphocyte migration index (LMI), and DNCB response in 66 sea-level residents, 45 temporary residents, and 24 natives at high altitude (3,692 m). An accentuated CMI, indicated by increase in PHA-blasts, increased lymphocyte migration index, and intense DNCB response, was present, despite a mild decrease in total leucocytes in temporary residents and in lymphocytes in natives at high altitude. While T-rosettes did not show any change in numbers, B-rosettes were increased in temporary residents, and natives at high altitude. A qualitative change had, therefore, occurred in lymphocytes at high altitude. CMI is equally augmented in temporary residents and natives at high altitude and prevails at a higher plane than at sea-level. Augmentation of CMI at high altitude, therefore, could be used as a therapeutic measure.     

Aspects of physical activity in high altitude natives

Altitude adaptation was measured at sea level and high altitudes in sea level adapted and high altitude adapted natives. Tests of work capacity as measured by O₂ consumption, pulse rate, and ventilation rate are reported. It is noted that comparing our results with those of other investigators is difficult due to variations in terminology and procedure. This suggests the necessity of more precise definitions in studies of the physical activity of high altitude natives.     

Carbonic anhydrase activity in the red blood cells of sea level and high altitude natives

Red blood cell carbonic anhydrase (CA) activity has not been studied in high altitude natives. Because CA is an intraerythocytic enzyme and high altitude natives are polycythemic, it is important to know if the activity of CA per red cell volume is different from that of their sea level counterparts. Blood was collected from healthy subjects living in Lima (150m) and from twelve subjects from Cerro de Pasco (4330m), and hematocrit and carbonic anhydrase activity were measured. As expected, the high altitude natives had significantly higher hematocrits than the sea level controls (p = 0.0002). No difference in the CA activity per milliliter of red cells was found between the two populations. There was no correlation between the hematocrit and CA activity.     

Developmental,genetic, and environmental components of aerobic capacity at high altitude

The aerobic capacity of 268 subjects (158 males and 110 females) was evaluated in La Paz, Bolivia situated at 3,750 m. The sample included 1) 39 high altitude rural natives (all male); 2) 67 high altitude urban natives (32 male, 35 female); 3) 69 Bolivians of foreign ancestry acclimatized to high altitude since birth (37 male, 32 female); 4) 50 Bolivians of foreign ancestry acclimatized to high altitude during growth (25 male, 25 female); and 5) 43 non-Bolivians of either European or North American ancestry acclimatized to high altitude during adulthood (25 male, 18 female). Data analyses indicate that 1) high altitude urban natives, acclimatized to high altitude since birth or during growth, attained higher aerobic capacity than subjects acclimatized to high altitude during adulthood; 2) age at arrival to high altitude is inversely related to maximum oxygen consumption (V?O² max) expressed in terms L/min or ml/min/kg of lean body mass, but not in terms of ml/min/kg of body weight; 3) among subjects acclimatized to high altitude during growth, approximately 25% of the variability in aerobic capacity can be explained by developmental factors; 4) as inferred from evaluations of skin color reflectance and sibling similarities, approximately 20 to 25% of the variability in aerobic capacity at high altitude can be explained by genetic factors; 5) except among the non-Bolivians acclimatized to high altitude during adulthood, the aerobic capacity of individuals with high occupational activity level is equal to the aerobic capacity of high altitude rural natives; and 6) the relationship between occupational activity level and aerobic capacity is much greater among subjects acclimatized to high altitude before the age of 10 years than afterwards. Together these data suggest that the attainment of normal aerobic capacity at high altitude is related to both developmental acclimatization and genetic factors but its expression is highly mediated by environmental factors, such as occupational activity level and body composition. © 1995 Wiley-Liss, Inc.     

Growth trends among male Bods of Ladakh--a high altitude population

A random sample of 274 native male Bods of Ladakh ranging in age from 11 to 19 years, living in Leh (Ladakh) at mean altitude of 3,514 metres, was studied. The pattern of growth of this sample was compared with the sea level Indians. The results indicate that: (1) The so called adolescent spurt is not well defined among Bod highlanders. (2) Bods grow faster than plain dwelling Indians and are taller and heavier at the age of 19 years. (3) The highlanders exhibit greater chest circumference than the lowland norms. This finding, similar to those among Andean natives, supports high altitude hypoxia's role in human growth and morphology.     

Adaptation and conservation of physiological systems in the evolution of human hypoxia tolerance

Analysis of human responses to hypobaric hypoxia in different lineages (lowlanders, Andean natives, Himalayan natives, and East Africans) indicates 'conservative' and 'adaptable' physiological characters involved in human responses to hypoxia. Conservative characters, derived by common descent, dominate and indeed define human physiology, but in five hypoxia response systems analyzed, we also found evidence for 'adaptable' characters at all levels of organization in all three high altitude lineages. Since Andeans and Himalayans have not shared common ancestry with East Africans for most of our species history, we suggest that their similar hypoxia physiology may represent the 'ancestral' condition for humans--an interpretation consistent with recent evidence indicating that our species evolved under 'colder, drier, and higher' conditions in East Africa where the phenotype would be simultaneously advantageous for endurance performance and for high altitude hypoxia. It is presumed that the phenotype was retained in low capacity form in highlanders and in higher capacity form in most lowland lineages (where it would be recognized by most physiologists as an endurance performance phenotype). Interestingly, it is easier for modern molecular evolution theory to account for the origin of 'adaptable' characters through positive selection than for conserved traits. Many conserved physiological systems are composed of so many gene products that it seems difficult to account for their unchanging state (for unchanging structure and function of hundreds of proteins linked in sequence to form the physiological system) by simple models of stabilizing selection.     

Longitudinal studies on serum proteins in man during two years of stay at an altitude of 4,000 m

The sequential changes in serum total protein concentration and in various electrophoretic fractions among lowlanders during two years of stay at high altitude (4,000 m) were determined and compared with that of high altitude natives (Ladakhis). The albumin to globulin ratio decreased during the early period of exposure to altitude among lowlanders and continued to remain at that level with minor fluctuations during the entire period of stay at altitude. Among the high altitude natives, a decreased albumin to globulin ratio with low serum protein concentration was observed. The composition of globulins separated electro-phoretically was of similar pattern in altitude natives and in lowlanders after a stay of two years at altitude. Among these subjects the-globulin was significantly higher as compared to values at sea level.     

Developmental components of resting ventilation among high- and low-altitude Andean children and adults

This paper evaluates the age-associated changes of resting ventilation of 115 high- and low-altitude Aymara subjects, of whom 61 were from the rural Aymara village of Ventilla situated at an average altitude of 4,200 m and 54 from the rural village of Caranavi situated at an average altitude of 900 m. Comparison of the age patterns of resting ventilation suggests the following conclusions: 1) the resting ventilation (ml/kg/min) of high-altitude natives is markedly higher than that of low-altitude natives; 2) the age decline of ventilation is similar in both lowlanders and highlanders, but the starting point and therefore the age decline are much higher at high altitude; 3) the resting ventilation that characterizes high-altitude Andean natives is developmentally expressed in the same manner as it is at low altitude; and 4) the resting ventilation (ml/kg/min) of Aymara high-altitude natives is between 40–80% lower than that of Tibetans. Am J Phys Anthropol 109:295–301, 1999. © 1999 Wiley-Liss, Inc.     

Body structure and respiratory efficiency among high altitude Himalayan populations

To understand the morphological and physiological variations among the temporary and permanent residents of high altitude, this study was undertaken at Leh, Ladakh. It is situated at 3500 m (11500 feet) above sea level, the mean barometric pressure was 500 tors and air temperature varied from 2 degrees C to 20 degrees C. The highland Tibetans showed broadest chest and most developed musculature closely followed by Ladakhi Bods. These high altude natives also displayed significantly higher value of vital capacity, forced vital capacity, and inspiratory capacity. The better respiratory efficiency observed among high altitude residents indicates higher degree of adaptation to high altitude hypoxia. Temporary residents were observed to be tallest and fattest with lower trunk fat predominance of all the four groups and showed narrowest chest and lower respiratory efficiency as compared to high altitude natives. The duration of stay at high altitude has clearly brought about a perceptible difference in body dimensions and respiratory functions of various groups of adult males studied at same altitude.     

Growth strategy,phylogeny and stoichiometry determine the allelopathic potential of native and non‐native plants

Secondary compounds can contribute to the success of non‐native plant species if they reduce damage by native herbivores or inhibit the growth of native plant competitors. However, there is opposing evidence on whether the secondary compounds of non‐native plant species are stronger than those of natives. This may be explained by other factors, besides plant origin, that affect the potential of plant secondary compounds. We tested how plant origin, phylogeny, growth strategy and stoichiometry affected the allelopathic potential of 34 aquatic plants. The allelopathic potential was quantified using bioassays with the cyanobacterium Dolichospermum flos‐aquae. The allelopathic potential showed a strong phylogenetic signal, but was similar for native and non‐native species. Growth strategy was important, and emergent plants had twice the allelopathic potential as compared to submerged plants. Furthermore, the allelopathic potential was positively correlated to the foliar carbon‐to‐phosphorus (C:P) and total phenolic content. We conclude that eudicot plant species with an emergent growth strategy and high plant C:P ratio exhibit a high allelopathic potential. Unless non‐native plant species match this profile, they generally have a similar allelopathic potential as natives.     

Evolutionary responses of natives to introduced species: what do introductions tell us about natural communities?

Biological invasions dramatically affect the distribution, abundance and reproduction of many native species. Because of these ecological effects, exotic species can also influence the evolution of natives exposed to novel interactions with invaders. Evolutionary changes in natives in response to selection from exotics are usually overlooked, yet common responses include altered anti-predator defenses, changes in the spectrum of resources and habitats used, and other adaptations that allow native populations to persist in invaded areas. Whether a native population is capable of responding evolutionarily to selection from invaders will depend on the demographic impact of the invader, the genetic architecture and genetic variability of the native population and potentially the history of previous invasions. In some cases, natives will fail to evolve or otherwise adapt, and local or global extinction will result. In other cases, adaptive change in natives may diminish impacts of invaders and potentially promote coexistence between invaders and natives. Here, we review the evidence for evolutionary responses of native species to novel community members. We also discuss how the effects of introduced species may differ from those caused by natural range expansions of native species. Notably, introduced species may come from remote biotas with no previous evolutionary history with the native community. In addition, the rate of addition of introduced species into communities is much greater than all but the most extreme cases of historical biotic exchange. Understanding the evolutionary component of exotic/native species interactions is critical to recognizing the long-term impacts of biological invasions, and to understanding the role of evolutionary processes in the assembly and dynamics of natural communities.     

Body water metabolism in high altitude natives during and after a stay at sea level

Body fluid compartments were studied in a group of high altitude natives after a stay of two months at sea level and during 12 days at an altitude of 3,500 m. Measurements of total body water and extracellular water were made on day 3 and 12 of reinduction to altitude, while plasma volume was measured on day 12 only. The intracellular water, blood volume and red cell mass were computed from the above parameters. Total body water and intracellular water decreased by 3.3% (P<0.001) and 5.0% (P<0.001) respectively by the 3rd day at altitude and did not change thereafter. Extracellular water increased progressively at altitude, but the increase was not significant. Blood volume and red cell mass increased significantly while plasma volume decreased at altitude. These data were compared with that of low landers. This study suggested body hypohydration on high altitude induction in low landers as well as in high altitude natives on reinduction.     

Local and landscape correlates of non‐native species invasion in restored wetlands

Vulnerability of natural communities to invasion by non‐native plants has been linked to factors such as recent disturbance and high resource availability, suggesting that recently restored habitats may be especially invasible. Because non‐native plants can interfere with restoration goals, monitoring programs should anticipate which sites are most susceptible to invasion and which species are likely to become problematic at a site. Restored sites of larger area and those with high rates of propagule input should have higher species richness of both natives and non‐natives, leading to a positive correlation between the two. However, in restored wetlands, urbanization, riparian landscape settings, and nitrogen enrichment likely favor non‐native relative to native species. We sampled 28 restored wetlands in Illinois, USA, modeled the responses of native richness, non‐native richness and non‐native cover to local and landscape predictors with linear regression, and modeled the presence/absence of 21 non‐native species with logistic regressions. Unexpectedly, native and non‐native richness were uncorrelated, suggesting different responses to environmental factors. Native richness declined with increasing available soil nitrogen and urbanization in the surrounding landscape. Non‐native richness, the richness of non‐natives relative to natives, and the likelihood of invasion by several individual invasive species decreased with increasing distance from the city of Chicago, likely in response to decreasing non‐native propagule pressure. Total cover of non‐natives, however, as well as cover by non‐native Phalaris arundinacea, increased with nitrogen availability. Our results indicate that although non‐native richness was better predicted by factors related to propagule pressure, non‐native species dominance was more closely related to local abiotic factors. Non‐native richness in restoration sites may be beyond the control of restoration practitioners, and furthermore, may be of limited relevance for conservation goals. In contrast, limiting the relative dominance of non‐natives should be a restoration priority and may be achievable through management of nutrient availability.     

Contrasting response of native and alien plant species richness to environmental energy and human impact along alpine elevation gradients

Aim We tested whether the species–energy and species–human relationships vary between native and both naturalized and casual alien species richness when other environmental variables had been taken into account. Location Trento Province, a region (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided into 156 contiguous (c. 37.5 km²) cells and ranging in elevation from 66 to 3769 m. Methods Data were separated into three subsets, representing richness of natives, naturalized aliens and casual aliens and separately related to temperature, human population and various environmental correlates of plant species diversity. We applied ordinary least squares and simultaneous autoregressive regressions to identify potential contrasting responses of the three plant status subsets and hierarchical partitioning to evaluate the relative importance of the predictor variables. Results Variation in alien plant species richness along the region was almost entirely explained by temperature and human population density. The relationships were positive but strongly curvilinear. Native species richness was less strongly related to either factor but was positively related to the presence of calcareous bedrock. Native species richness had a decelerating positive relationship with temperature (R²= 55%), whereas naturalized and casual aliens had a positive accelerating relationship explaining 86% and 62% of the variation in richness, respectively. Native species richness had a positive decelerating relationship with population density (R²= 42%), whilst both alien subsets had a positive accelerating relationship. Main conclusions Alien species richness was higher in areas with the most rich and diverse assemblages of native species. Areas at high altitudes are not especially prone to alien invasion due to energy constraints, low propagule pressure and disturbance, even considering a potential increased in temperature. Thus, if we consider future environmental change, we should expect a stronger response of aliens than natives in the currently warm, urbanized, low‐altitude areas than in cold, high‐altitude areas where human population density is low.     

Neighbour tolerance,not suppression,provides competitive advantage to non‐native plants

High competitive ability has often been invoked as a key determinant of invasion success and ecological impacts of non‐native plants. Yet our understanding of the strategies that non‐natives use to gain competitive dominance remains limited. Particularly, it remains unknown whether the two non‐mutually exclusive competitive strategies, neighbour suppression and neighbour tolerance, are equally important for the competitive advantage of non‐native plants. Here, we analyse data from 192 peer‐reviewed studies on pairwise plant competition within a Bayesian multilevel meta‐analytic framework and show that non‐native plants outperform their native counterparts due to high tolerance of competition, as opposed to strong suppressive ability. Competitive tolerance ability of non‐native plants was driven by neighbour's origin and was expressed in response to a heterospecific native but not heterospecific non‐native neighbour. In contrast to natives, non‐native species were not more suppressed by hetero‐ vs. conspecific neighbours, which was partially due to higher intensity of intraspecific competition among non‐natives. Heterogeneity in the data was primarily associated with methodological differences among studies and not with phylogenetic relatedness among species. Altogether, our synthesis demonstrates that non‐native plants are competitively distinct from native plants and challenges the common notion that neighbour suppression is the primary strategy for plant invasion success.     

Association of high-altitude systemic hypertension with the deletion allele-of the angiotensin-converting enzyme (ACE) gene

People who visit high-altitude areas are exposed to a stressful environment and a good percentage of them suffer from high-altitude-induced diseases, including systemic hypertension. Identification of genetic markers for high-altitude-induced diseases would help to reduce the rate of morbidity/mortality from such diseases. The development of systemic hypertension on exposure to high altitude (3,500 m) for 30 days in otherwise normotensive natives of low-altitudes was investigated. The angiotensin-converting enzyme (ACE) insertion/deletion (I/D) genotypes and renin-angiotensin-aldosterone system were simultaneously studied. In the hypertensives during their stay at high altitude, the ACE D allele frequency was significantly higher than in the normotensives (0.67 versus 0.32 chi(2)(1) = 10.6, P < 0.05). In the normotensives during their stay at high altitude, there was no significant increase in plasma aldosterone levels despite increased plasma renin activity. Results of the present study suggest that environmental changes and pre-existing genetic factors, namely the ACE D allele, might be two of the factors predisposing natives of low altitudes to systemic hypertension, a polygenic disease, at high altitude.