Organisms at high altitude

Source: Wikipedia, the free encyclopedia.
An Alpine chough in flight at 3,901 m (12,799 ft)

Organisms can live at

vicunas, llamas, mountain goats, etc.) and certain birds are known to have completely adapted to high-altitude environments.[1]

Human populations such as some

Ethiopians live in the otherwise uninhabitable high mountains of the Himalayas, Andes and Ethiopian Highlands respectively. The adaptation of humans to high altitude is an example of natural selection in action.[2]

High-altitude adaptations provide examples of convergent evolution, with adaptations occurring simultaneously on three continents. Tibetan humans and Tibetan domestic dogs share a genetic mutation in EPAS1, but it has not been seen in Andean humans.[3]

Invertebrates

Tardigrades live over the entire world, including the high Himalayas.[4] Tardigrades are also able to survive temperatures of close to absolute zero (−273 °C (−459 °F)),[5] temperatures as high as 151 °C (304 °F), radiation that would kill other animals,[6] and almost a decade without water.[7] Since 2007, tardigrades have also returned alive from studies in which they have been exposed to the vacuum of outer space in low Earth orbit.[8][9]

Other invertebrates with high-altitude habitats are Euophrys omnisuperstes, a spider that lives in the Himalaya range at altitudes of up to 6,699 m (21,978 ft);[10] it feeds on stray insects that are blown up the mountain by the wind.[11] The springtail Hypogastrura nivicola (one of several insects called snow fleas) also lives in the Himalayas. It is active in the dead of winter, its blood containing a compound similar to antifreeze. Some allow themselves to become dehydrated instead, preventing the formation of ice crystals within their body.[12]

Insects can fly and kite at very high altitude.

Bumble bees were discovered on Mount Everest at more than 5,600 m (18,400 ft) above sea level.[14] In subsequent tests, bumblebees were still able to fly in a flight chamber which recreated the thinner air of 9,000 m (30,000 ft).[15]

Ballooning is a term used for the mechanical kiting[16][17] that many spiders, especially small species such as Erigone atra,[18] as well as certain mites and some caterpillars use to disperse through the air. Some spiders have been detected in atmospheric data balloons collecting air samples at slightly less than 5 km (16000 ft) above sea level.[19] It is the most common way for spiders to pioneer isolated islands and mountaintops.[20][21]

Fish

Lake Qinghai
at 3,205 m (10,515 ft)

Fish at high altitudes have a lower metabolic rate, as has been shown in highland westslope cutthroat trout when compared to introduced lowland rainbow trout in the Oldman River basin.[22] There is also a general trend of smaller body sizes and lower species richness at high altitudes observed in aquatic invertebrates, likely due to lower oxygen partial pressures.[23][24][25] These factors may decrease productivity in high altitude habitats, meaning there will be less energy available for consumption, growth, and activity, which provides an advantage to fish with lower metabolic demands.[22]

The

hypoxia-inducible factor 1 (HIF-1).[27]
It is unclear whether this is a common characteristic in other high altitude dwelling fish or if gill remodelling and HIF-1 use for cold adaptation are limited to carp.

Mammals

The Himalayan pika lives at altitudes up to 4,200 m (13,800 ft)[28]

herbivores of the Himalayas such as the Himalayan tahr, markhor and chamois are of particular interest because of their ecological versatility and tolerance.[32]

Rodents

A number of

carbohydrates for small bursts of energy.[35]

Other physiological changes that occur in rodents at high altitude include increased

breathing rate[36] and altered morphology of the lungs and heart, allowing more efficient gas exchange and delivery. Lungs of high-altitude mice are larger, with more capillaries,[37] and their hearts have a heavier right ventricle (the latter applies to rats too),[38][39]
which pumps blood to the lungs.

At high altitudes, some rodents even shift their thermal neutral zone so they may maintain normal basal metabolic rate at colder temperatures.[40]

The deer mouse

The deer mouse (

Phyllotis limatus). This shows that highland mice have evolved a metabolic process to economise oxygen usage for physical activities in the hypoxic conditions.[46]

Yaks

Domestic yak at Yamdrok Lake
Main article: Yak

Among

sensory perception and energy metabolism were identified.[49] In addition, researchers also found an enrichment of protein domains related to the extracellular environment and hypoxic stress that had undergone positive selection and rapid evolution. For example, they found three genes that may play important roles in regulating the bodyʼs response to hypoxia, and five genes that were related to the optimisation of the energy from the food scarcity in the extreme plateau. One gene known to be involved in regulating response to low oxygen levels, ADAM17, is also found in human Tibetan highlanders.[50][51]

Humans

Main articles: High-altitude adaptation in humans and Effects of high altitude on humans
A Sherpa family

Over 81 million people live permanently at high altitudes (>2,500 m (8,200 ft))[52] in North, Central and South America, East Africa, and Asia, and have flourished for millennia in the exceptionally high mountains, without any apparent complications.[53] For average human populations, a brief stay at these places can risk mountain sickness.[54] For the native highlanders, there are no adverse effects to staying at high altitude.

The physiological and genetic adaptations in native highlanders involve modification in the oxygen transport system of the blood, especially molecular changes in the structure and functions of hemoglobin, a protein for carrying oxygen in the body.[53][55] This is to compensate for the low oxygen environment. This adaptation is associated with developmental patterns such as high birth weight, increased lung volumes, increased breathing, and higher resting metabolism.[56][57]

The

DNA damage stimulus and DNA repair (such as RAD51, RAD52, and MRE11A), which are related to the adaptive traits of high infant birth weight and darker skin tone and, are most likely due to recent local adaptation.[61]

Among the Andeans, there are no significant associations between EPAS1 or EGLN1 and hemoglobin concentration, indicating variation in the pattern of molecular adaptation.

HIF genetic functions.[65]

The EPAS1 mutation in the Tibetan population has been linked to Denisovan-related populations.[66] The Tibetan haplotype is more similar to the Denisovan haplotype than any modern human haplotype. This mutation is seen at a high frequency in the Tibetan population, a low frequency in the Han population and is otherwise only seen in a sequenced Denisovan individual. This mutation must have been present before the Han and Tibetan populations diverged 2750 years ago.[66]

Birds

Rüppell's vulture can fly up to 11.2 km (7.0 mi) above sea level
See also: List of birds by flight heights

Birds have been especially successful at living at high altitudes.

capillaries and small muscle fibres (which increases surface-area-to-volume ratio).[71] These two features facilitate oxygen diffusion from the blood to muscle, allowing flight to be sustained during environmental hypoxia. Birds' hearts and brains, which are very sensitive to arterial hypoxia, are more vascularized compared to those of mammals.[72] The bar-headed goose (Anser indicus) is an iconic high-flyer that surmounts the Himalayas during migration,[73] and serves as a model system for derived physiological adaptations for high-altitude flight. Rüppell's vultures, whooper swans, alpine chough, and common cranes
all have flown more than 8 km (26,000 ft) above sea level.

Adaptation to high altitude has fascinated

jays and geese.[29][31][74]
The Andes is quite rich in bird diversity. The
diuca-finches are also found in the highlands.[75][76]

Cinnamon teal

Male cinnamon teal

Evidence for adaptation is best investigated among the Andean birds. The

substitution at position 9 of the protein, with asparagine present almost exclusively within the low-elevation species, and serine in the high-elevation species. This implies important functional consequences for oxygen affinity.[77] In addition, there is strong divergence in body size in the Andes and adjacent lowlands. These changes have shaped distinct morphological and genetic divergence within South American cinnamon teal populations.[78]

Ground tits

In 2013, the molecular mechanism of high-altitude adaptation was elucidated in the Tibetan ground tit (

ADRBK1 and HSD17B7, which are involved in the adrenaline response and steroid hormone biosynthesis. Thus, the strengthened hormonal system is an adaptation strategy of this bird.[79]

Other animals

amphibians in the Tibetan highlands.[74] Gloydius himalayanus is perhaps the geographically highest living snake in the world, living at as high as 4,900 m (16,100 ft) in the Himalayas.[80] Another notable species is the Himalayan jumping spider, which can live at over 6,500 m (21,300 ft) of elevation.[29]

Plants

Main article:
Alpine plants
Cushion plant Donatia novae-zelandiae, Tasmania

Many different plant species live in the high-altitude environment. These include

perennial grasses, sedges, forbs, cushion plants, mosses, and lichens.[81] High-altitude plants must adapt to the harsh conditions of their environment, which include low temperatures, dryness, ultraviolet radiation, and a short growing season. Trees cannot grow at high altitude, because of cold temperature or lack of available moisture.[82]: 51  The lack of trees causes an ecotone, or boundary, that is obvious to observers. This boundary is known as the tree line
.

The highest-altitude plant species is a

Arenaria bryophylla is the highest flowering plant in the world, occurring as high as 6,180 m (20,280 ft).[84]

See also

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