Wikipedia

Organisms at high altitude

Organisms can live athigh altitude,either on land, in water, or while flying. Decreased oxygen availability and decreased temperature make life at such altitudes challenging, though many species have been successfullyadaptedvia considerablephysiologicalchanges. As opposed to short-termacclimatisation(immediate physiological response to changing environment), high-altitudeadaptationmeans irreversible,evolved physiological responsesto high-altitude environments, associated with heritablebehaviouralandgenetic changes.Among vertebrates, only few mammals (such asyaks,ibexes,Tibetan gazelles,vicunas,llamas,mountain goats,etc.) and certainbirdsare known to have completely adapted to high-altitude environments.[1]

AnAlpine choughin flight at 3,900 m (12,800 ft)

Human populations such as someTibetans,South AmericansandEthiopianslive in the otherwise uninhabitable high mountains of theHimalayas,AndesandEthiopian Highlandsrespectively. The adaptation of humans to high altitude is an example ofnatural selectionin action.[2]

High-altitude adaptations provide examples ofconvergent evolution,with adaptations occurring simultaneously on three continents. Tibetan humans and Tibetan domestic dogs share a genetic mutation inEPAS1,but it has not been seen in Andean humans.[3]

Invertebrates

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Tardigradeslive over the entire world, including the highHimalayas.[4]Tardigrades are also able to survive temperatures of close toabsolute zero(−273 °C or −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 areEuophrys omnisuperstes,a spider that lives in the Himalaya range at altitudes of up to 6,700 m (22,000 ft);[10]it feeds on stray insects that are blown up the mountain by the wind.[11]ThespringtailHypogastrura 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 toantifreeze.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.Fliesare common in the Himalayas up to 6,300 m (20,700 ft).[13]Bumble beeswere discovered onMount Everestat 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]

Ballooningis a term used for the mechanical kiting[16][17]that manyspiders,especially small species such asErigone atra,[18]as well as certainmitesand somecaterpillarsuse to disperse through the air. Some spiders have been detected in atmospheric data balloons collecting air samples at slightly less than 5 km (16,000 ft) above sea level.[19]It is the most common way for spiders to pioneer isolated islands and mountaintops.[20][21]

Fish

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Naked carp inLake Qinghaiat 3,205 m (10,515 ft)

Fish at high altitudes have a lower metabolic rate, as has been shown in highlandwestslope cutthroat troutwhen compared to introduced lowlandrainbow troutin theOldman Riverbasin.[22]There is also a general trend of smaller body sizes and lowerspecies richnessat high altitudes observed in aquatic invertebrates, likely due to lower oxygen partial pressures.[23][24][25]These factors may decreaseproductivityin 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]

Thenaked carpfromLake Qinghai,like other members of thecarpfamily, can usegill remodellingto increase oxygen uptake inhypoxic environments.[26]The response of naked carp to cold and low-oxygen conditions seem to be at least partly mediated byhypoxia-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

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TheHimalayan pikalives at altitudes up to 4,200 m (13,800 ft)[28]

Mammalsare also known to reside at high altitude and exhibit a striking number of adaptations in terms ofmorphology,physiologyandbehaviour.TheTibetan Plateauhas very few mammalian species, ranging fromwolf,kiang(Tibetan wild ass),goas,chiru(Tibetan antelope),wild yak,snow leopard,Tibetan sand fox,ibex,gazelle,Himalayan brown bearandwater buffalo.[29][30][31]These mammals can be broadly categorised based on their adaptability in high altitude into two broad groups, namelyeurybarcandstenobarc.Those that can survive a wide range of high-altitude regions areeurybarcand include yak, ibex,Tibetan gazelleof the Himalayas andvicuñasllamasof the Andes. Stenobarc animals are those with lesser ability to endure a range of differences in altitude, such asrabbits,mountain goats,sheep,andcats.Amongdomesticated animals,yaks are perhaps the highest dwelling animals. Thewildherbivoresof the Himalayas such as theHimalayan tahr,markhorandchamoisare of particular interest because of their ecological versatility and tolerance.[32]

Rodents

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A number ofrodentslive at high altitude, includingdeer mice,guinea pigs,andrats.Several mechanisms help them survive these harsh conditions, including alteredgeneticsof thehemoglobingene in guinea pigs and deer mice.[33][34]Deer mice use a high percentage of fats as metabolic fuel to retaincarbohydratesfor small bursts of energy.[35]

Other physiological changes that occur in rodents at high altitude include increasedbreathing rate[36]and altered morphology of the lungs and heart, allowing more efficientgas exchangeand 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 theirthermal neutral zoneso they may maintain normalbasal metabolic rateat colder temperatures.[40]

The deer mouse

The deer mouse (Peromyscus maniculatus) is the best studied species, other than humans, in terms of high-altitude adaptation.[1]The deer mice native to Andes highlands (up to 3,000 m (9,800 ft)) are found to have relatively low hemoglobin content.[41]Measurement of food intake,gutmass, andcardiopulmonary organmass indicated proportional increases in mice living at high altitudes, which in turn show that life at high altitudes demands higher levels of energy.[42]Variations in theglobingenes (αandβ-globin) seem to be the basis for increased oxygen-affinity of the hemoglobin and faster transport of oxygen.[43][44]Structural comparisons show that in contrast to normal hemoglobin, the deer mouse hemoglobin lacks thehydrogen bondbetweenα1Trp14in the Ahelixand α1Thr67 in the E helix owing to theThr67Alasubstitution, and there is a unique hydrogen bond at the α1β1 interface between residuesα1Cys34andβ1Ser128.[45]The Peruvian native species of mice (Phyllotis andiumandPhyllotis xanthopygus) have adapted to the high Andes by using proportionately morecarbohydratesand have higher oxidative capacities ofcardiac musclescompared to closely related native species residing at low-altitudes (100–300 m (330–980 ft)), (Phyllotis amicusandPhyllotis limatus). This shows that highland mice have evolved a metabolic process to economise oxygen usage for physical activities in the hypoxic conditions.[46]

Yaks

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Domestic yak atYamdrok Lake
Main article:Yak

Amongdomesticated animals,yaks (Bos grunniens) are the highest dwelling animals of the world, living at 3,000–5,000 m (9,800–16,400 ft). The yak is the most important domesticated animal for Tibet highlanders inQinghai ProvinceofChina,as the primary source ofmilk,meatandfertilizer.Unlike other yak orcattlespecies, which suffer from hypoxia in the Tibetan Plateau, the Tibetan domestic yaks thrive only at high altitude, and not in lowlands. Their physiology is well-adapted to high altitudes, with proportionately larger lungs and heart than other cattle, as well as greater capacity for transporting oxygen through their blood.[47]In yaks,hypoxia-inducible factor1 (HIF-1) has high expression in thebrain,lungandkidney,showing that it plays an important role in the adaptation to low oxygen environment.[48]On 1 July 2012 the complete genomic sequence and analyses of a female domestic yak was announced, providing important insights into understanding mammaliandivergenceand adaptation at high altitude. Distinct gene expansions related tosensory perceptionand 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

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Main articles:High-altitude adaptation in humansandEffects of high altitude on humans
ASherpafamily

Over 81 million people live permanently at highaltitudes(>2,500 m or 8,200 ft)[52]inNorth,CentralandSouth America,East Africa,andAsia,and have flourished formillenniain the exceptionally high mountains, without any apparent complications.[53]For average human populations, a brief stay at these places can riskmountain sickness.[54]For the native highlanders, there are no adverse effects to staying at high altitude.

The physiological and geneticadaptationsin native highlanders involve modification in theoxygen transport system of the blood,especiallymolecular changesin the structure and functions ofhemoglobin,a protein for carrying oxygen in the body.[53][55]This is to compensate for thelow oxygen environment.This adaptation is associated with developmental patterns such as highbirth weight,increasedlung volumes,increasedbreathing,and higherresting metabolism.[56][57]

Thegenomeof Tibetans provided the first clue to themolecular evolutionof high-altitude adaptation in 2010.[58]Genes such asEPAS1,PPARAandEGLN1are found to have significantmolecular changesamong the Tibetans, and the genes are involved inhemoglobin production.[59]These genes function in concert with transcription factors,hypoxia inducible factors(HIF), which in turn are central mediators ofred blood cell productionin response to oxygen metabolism.[60]Further, the Tibetans are enriched for genes in the disease class of human reproduction (such as genes from theDAZ,BPY2,CDY,andHLA-DQandHLA-DRgene clusters) and biological process categories of response toDNA damagestimulus andDNA repair(such asRAD51,RAD52,andMRE11A), which are related to the adaptive traits of high infant birth weight anddarker skin toneand, are most likely due to recent local adaptation.[61]

Among the Andeans, there are no significant associations betweenEPAS1orEGLN1and hemoglobin concentration, indicating variation in the pattern of molecular adaptation.[62]However,EGLN1appears to be the principal signature of evolution, as it shows evidence of positive selection in both Tibetans and Andeans.[63]The adaptive mechanism is different among the Ethiopian highlanders. Genomic analysis of two ethnic groups,AmharaandOromo,revealed that gene variations associated with hemoglobin differences among Tibetans or other variants at the samegene locationdo not influence the adaptation in Ethiopians.[64]Instead, several other genes appear to be involved in Ethiopians, includingCBARA1,VAV3,ARNT2andTHRB,which are known to play a role inHIFgenetic functions.[65]

The EPAS1 mutation in the Tibetan population has been linked toDenisovan-related populations.[66]The Tibetanhaplotypeis 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

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Rüppell's vulturecan 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.[67]In general, birds have physiological features that are advantageous for high-altitude flight. Therespiratory system of birdsmoves oxygen across the pulmonary surface during both inhalation and exhalation, making it more efficient than that of mammals.[68]In addition, the air circulates in one direction through theparabronchiolesin the lungs. Parabronchioles are oriented perpendicularly to thepulmonary arteries,forming across-current gas exchanger.This arrangement allows for more oxygen to be extracted compared to mammalianconcurrent gas exchange;as oxygen diffuses down its concentration gradient and the air gradually becomes more deoxygenated, the pulmonary arteries are still able to extract oxygen.[69][page needed]Birds also have a high capacity for oxygen delivery to the tissues because they have larger hearts and cardiacstroke volumecompared to mammals of similar body size.[70]Additionally, they have increased vascularization in their flight muscle due to increased branching of thecapillariesand small muscle fibres (which increasessurface-area-to-volumeratio).[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]Thebar-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,andcommon cranesall have flown more than 8 km (26,000 ft) above sea level.

Adaptation to high altitude has fascinatedornithologistsfor decades, but only a small proportion of high-altitude species have been studied. In Tibet, few birds are found (28endemic species), includingcranes,vultures,hawks,jaysandgeese.[29][31][74] The Andes is quite rich in bird diversity. TheAndean condor,the largest bird of its kind in theWestern Hemisphere,occurs throughout much of the Andes but generally in very low densities; species oftinamous(notably members of the genusNothoprocta),Andean goose,giant coot,Andean flicker,diademed sandpiper-plover,mountain parakeet,miners,sierra-finchesanddiuca-finchesare also found in the highlands.[75][76]

Cinnamon teal

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Male cinnamon teal

Evidence for adaptation is best investigated among the Andean birds. Thewater fowlsand cinnamon teal (Anas cyanoptera) are found to have undergone significantmolecular modifications.It is now known that the α-hemoglobin subunit gene is highly structured between elevations among cinnamon teal populations, which involves almost entirely a single non-synonymousamino acidsubstitutionat position 9 of theprotein,withasparaginepresent almost exclusively within the low-elevation species, andserinein 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

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In 2013, the molecular mechanism of high-altitude adaptation was elucidated in the Tibetan ground tit (Pseudopodoces humilis) using a draft genome sequence. Gene family expansion and positively selected gene analysis revealed genes that were related to cardiac function in the ground tit. Some of the genes identified to have positive selection includeADRBK1andHSD17B7,which are involved in theadrenalineresponse andsteroid hormone biosynthesis.Thus, the strengthenedhormonal systemis an adaptation strategy of this bird.[79]

Other animals

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AlpineTibet hosts a limited diversity of animal species, among whichsnakesare common. There are only two endemicreptilesand ten endemicamphibiansin the Tibetan highlands.[74]Gloydius himalayanusis 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 theHimalayan jumping spider,which can live at over 6,500 m (21,300 ft) of elevation.[29]

Plants

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Main article:Alpine plants
Cushion plantDonatia novae-zelandiae,Tasmania

Many different plant species live in the high-altitude environment. These includeperennial grasses,sedges,forbs,cushion plants,mosses,andlichens.[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 anecotone,or boundary, that is obvious to observers. This boundary is known as thetree line.

The highest-altitude plant species is amossthat grows at 6,480 m (21,260 ft) onMount Everest.[83]The sandwortArenaria bryophyllais the highest flowering plant in the world, occurring as high as 6,180 m (20,280 ft).[84]

See also

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