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• W3153073103 abstract "The genomics of adaptation to deserts is a rapidly growing research field that provides examples of adaptation over different timescales and of vastly different organisms facing shared challenges.Mammals inhabiting deserts show remarkable adaptive traits that have evolved repeatedly and independently in different species across the globe and in response to similar selective pressures of extreme temperatures, aridity, and water and food deprivation.Genomic studies have shown that there are shared patterns of adaptation at the genomic level involving fat metabolism and insulin signaling, as well as arachidonic acid metabolism.Understanding the mechanisms by which species have successfully adapted to the physical and climatic challenges of deserts is important for evaluating the possibility of evolutionary rescue of species currently challenged by increased desertification. Deserts are among the harshest environments on Earth. The multiple ages of different deserts and their global distribution provide a unique opportunity to study repeated adaptation at different timescales. Here, we summarize recent genomic research on the genetic mechanisms underlying desert adaptations in mammals. Several studies on different desert mammals show large overlap in functional classes of genes and pathways, consistent with the complexity and variety of phenotypes associated with desert adaptation to water and food scarcity and extreme temperatures. However, studies of desert adaptation are also challenged by a lack of accurate genotype–phenotype–environment maps. We encourage development of systems that facilitate functional analyses, but also acknowledge the need for more studies on a wider variety of desert mammals. Deserts are among the harshest environments on Earth. The multiple ages of different deserts and their global distribution provide a unique opportunity to study repeated adaptation at different timescales. Here, we summarize recent genomic research on the genetic mechanisms underlying desert adaptations in mammals. Several studies on different desert mammals show large overlap in functional classes of genes and pathways, consistent with the complexity and variety of phenotypes associated with desert adaptation to water and food scarcity and extreme temperatures. However, studies of desert adaptation are also challenged by a lack of accurate genotype–phenotype–environment maps. We encourage development of systems that facilitate functional analyses, but also acknowledge the need for more studies on a wider variety of desert mammals. Deserts (see Glossary) are the driest environments on the planet and cover at least 33% of the land surface on Earth [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar]. Although mainly characterized by aridity and water scarcity, deserts also experience daily and annual extreme thermal amplitudes, and intense UV radiation [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar]. Deserts have long been seen as natural laboratories for investigating how biological design is challenged by different aspects of the environment, and how organisms have adapted to these challenges [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar,2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. They also offer unique opportunities to study convergent evolution at distinct points in time and space, given their well-documented geological age and diverse geographical origins [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar,2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. The main challenges to life in deserts are maintaining body temperature and preserving water [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. This is particularly difficult for species that rely heavily on evaporative water loss mechanisms (e.g., sweating, panting, and salivation) for dissipating heat [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,3.Riddell E.A. et al.Exposure to climate change drives stability or collapse of desert mammal and bird communities.Science. 2021; 371: 633-638Crossref PubMed Scopus (40) Google Scholar]. Exposure to heat stress combined with water shortage in hot deserts triggers several pathophysiological processes resulting in heat exhaustion, heat stroke, and kidney dysfunction [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar,4.Glaser J. et al.Climate change and the emergent epidemic of CKD from heat stress in rural communities: the case for heat stress nephropathy.Clin. J. Am. Soc. Nephrol. 2016; 11: 1472-1483Crossref PubMed Scopus (222) Google Scholar,5.Johnson R.J. et al.Metabolic and kidney diseases in the setting of climate change, water shortage, and survival factors.J. Am. Soc. Nephrol. 2016; 27: 2247-2256Crossref PubMed Scopus (49) Google Scholar]. Exposure to extreme low temperatures in cold-arid deserts, or at night/winter in hot deserts, can lead to hypothermia [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. The organismal responses required to survive these extreme conditions for multiple generations are usually outside the range of the plastic responses of most non-desert species. Thus, it is generally presumed that desert-adapted species likely experienced strong selection acting on various complex phenotypes related to metabolism and water retention [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar,2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar, 7.Rocha J.L. et al.Convergent evolution of increased urine-concentrating ability in desert mammals.Mamm. Rev. 2021; (Published online March 1, 2021. https://doi.org/10.1111/mam.12244)Crossref PubMed Scopus (2) Google Scholar, 8.Williams J.B. Tieleman B.I. Physiological adaptation in desert birds.Bioscience. 2005; 55: 416-425Crossref Scopus (110) Google Scholar]. While desert adaptation has been subject to much research in evolutionary ecology, there have been relatively few investigations on the underlying genetic basis of desert adaptive traits. However, newly emerging research on desert mammalian case studies is contributing to our understanding of the role of past climatic processes, such as desertification, in driving adaptation. In this review, we focus on this nascent research field. We review different methods and synthesize current genomic work in mammalian systems. We then explore the limits and future potential of using genomics to address desert adaptation and provide explicit recommendations for future research. The recurrence of diverse complex adaptive phenotypes across diverse desert organisms (Box 1 for examples) leads to a fundamental interest in their underlying genetic basis. Enabled by increasingly cost-effective sequencing technologies and computational resources for data analysis, researchers may rely on numerous methods that were designed to identify regions associated with selection, the footprints of adaptation, from large genomic data sets [9.Stern A.J. Nielsen R. Detecting natural selection.in: Balding D.J. Handbook of Statistical Genomics. Wiley, 2019: 397-340Crossref Scopus (5) Google Scholar,10.Nielsen R. Molecular signatures of natural selection.Annu. Rev. Genet. 2005; 39: 197-218Crossref PubMed Scopus (1095) Google Scholar]. The choice of the sequencing and analytical approach should be guided by knowledge of the ecological and evolutionary context of the species, and by the possibility of biological sampling infrastructures in remote arid regions.Box 1Endurer–Evader–Evaporator ConceptAiming to summarize and categorize decades of classic literature on desert adaptations in species occupying distinct ecological niches, Willmer et al. [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar] proposed a classification system relative to extreme temperatures and body size. Animals that can evade extreme temperatures through behavior and are physiologically adapted to minimize water loss are called ‘evaders’. Mammalian evaders include small rodents that can avoid overheating by being nocturnal and hiding in burrows during the day, and are less reliant on evaporative cooling [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. Evaders can additionally minimize water loss by concentrating highly hyperosmotic urine with low volume [7.Rocha J.L. et al.Convergent evolution of increased urine-concentrating ability in desert mammals.Mamm. Rev. 2021; (Published online March 1, 2021. https://doi.org/10.1111/mam.12244)Crossref PubMed Scopus (2) Google Scholar] and utilize water derived from fat metabolism (metabolic water) well beyond the abilities of non-desert counterparts (Table I for examples) [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar].Animals that cannot shelter behaviorally and instead rely on morphology and physiology to minimize water loss and withstand heat are called ‘endurers’ (Table I) [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. Endurers include large ungulates and marsupials able to store heat without increasing their body temperature [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar]. For example, Dromedary camels (Camelus dromedarius) can endure temperatures exceeding 42°C, and water losses >25% of their total body weight, which are both fatal to non-desert mammals [41.Ali A. et al.From desert to medicine: a review of camel genomics and therapeutic products.Front. Genet. 2019; 10: 1-20Crossref PubMed Scopus (37) Google Scholar]. Some endurers also have lower respiratory water losses [6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar], and can store water in their rumen, gut, or intestines for water retention [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar]. It has also been proposed that Arabian Oryx (Oryx leucocoryx ) use water derived from fat metabolism and that this may account for 24% of their overall water needs [5.Johnson R.J. et al.Metabolic and kidney diseases in the setting of climate change, water shortage, and survival factors.J. Am. Soc. Nephrol. 2016; 27: 2247-2256Crossref PubMed Scopus (49) Google Scholar]. Endurers may also have enlarged body tissues that can store localized fat (e.g., fat tails in sheep, Ovis aries, and fat-filled humps of camels) to act as energy reserves during starvation [6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar,83.Atti N. et al.Performance of the fat-tailed Barbarine sheep in its environment: adaptive capacity to alternation of underfeeding and re-feeding periods. A review.Anim. Res. 2004; 53: 165-176Crossref Scopus (70) Google Scholar].A third category was additionally proposed for medium-sized animals that are not able to avoid extreme conditions as efficiently as evaders nor withstand heat as efficiently as endurers. These are called ‘evaporators’ and comprise lagomorphs, some marsupials, and medium-sized carnivores (Table I) [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. Evaporator species have enlarged body extremities (e.g., large ears) to dissipate heat by conduction and use denning and nocturnal behavior to avoid extreme heat. They also have significantly low mass-adjusted nonrenal water loss [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar,2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar] and can minimize renal water loss by concentrating highly hyperosmotic urine significantly above the levels seen in non-desert counterparts [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,7.Rocha J.L. et al.Convergent evolution of increased urine-concentrating ability in desert mammals.Mamm. Rev. 2021; (Published online March 1, 2021. https://doi.org/10.1111/mam.12244)Crossref PubMed Scopus (2) Google Scholar]. However, some authors have criticized this categorization due to overlap between the categories and have proposed other frameworks to synthesize desert mammalian phenotypic adaptations (e.g., [6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar]).Table IClassic Physiological Systems Allowing Mammalian Survival in DesertsaThe phenotypes listed may themselves be adaptive or may be part of a correlated plastic response ([6,7,41,52,65,84–87] for more examples and [6,65,66,85] for detailed reviews)., bSuperscript codes and numbers designate: En – Endurers (1, Dromedary camel, Camelus dromedarius; 2, Arabian oryx, Oryx leucoryx; 3, Black Bedouin goat, Capra aegagrus hircus; 4, Awassi fat-tailed sheep, Ovis aries; 5, Marwari sheep; 6, Desert bighorn sheep, Ovis canadensis nelsoni; 7, domestic Bactrian camel, Camelus bactrianus; 7*, wild Bactrian camel, Camelus ferus; 8, Ethiopian Somali goat, Capra aegagrus hircus; 9, Common wallaroo, Osphranter robustus; 10, Gemsbok, Oryx gazella; 11, Arabian sand gazelle, Gazella marica); Ev – Evaders (1, Golden spiny mouse, Acomys russatus; 2, Merriam's kangaroo rat, Dipodomys merriami; 3, Tarabul's gerbil, Gerbillus tarabuli; 4, Cairo spiny mouse, Acomys cahirinus; 5, Cactus mouse, Peromyscus eremicus, 6, Spinifex hopping mouse, Notomys alexis; 7, Sonoran desert mice, Mus musculus; 8, Mongolian gerbil, Meriones unguiculatus; 9, Fawn hopping mouse, Notomys cervinus; 10, Degu, Octodon degus; 11, Agile kangaroo rat, Dipodomys agilis; 12, Desert pocket mouse, Chaetodipus penicillatus; 13, Namib dune gerbil, Gerbillurus tytonis; 14, Fat sand rat, Psammomys obesus); Evap – Evaporators (1, Fennec fox, Vulpes zerda; 2, Rueppell’s fox, Vulpes rueppellii; 3, Kit fox, Vulpes macrotis; 4, Cape hare, Lepus capensis; 5, Tammar wallaby, Macropus eugenii; 6, Springhare, Pedetes capensis); H, Aboriginal Australians, Homo sapiens.Environmental stressorUpstream phenotypeDownstream phenotypeHeat + waterDecreased thyroxinEn (4,5), Ev (2,5), HLow metabolism or energy expendituresEn (1,2,9), Ev (1,2,8), Evap (1), HLow metabolism/energy expendituresReduced nonrenal water lossEn (1,2,10), Ev (2,8), Evap (1,2), H, low respiration rateEn (1,2), Ev (6), Evap (1), torporEv (1,5)ColdNonshivering thermogenesisEv (1,8)Reduced shiveringHHeatReduced aldosteroneEn (1)Low cardiac rate and low blood pressureEn (1)WaterIncreased vasopressinEn (1,8), Ev (1,6,10); Increased aldosteroneEn (1,3,4), Ev (3,4)Increased urine osmolality/Increased water reabsorption from the kidneyEn (1,2), Ev, Evap (1,4); decreased urine productionEv (1,5,7)Reduced aldosteroneEn (1)Increased urine sodium excretionEn (1)Low glomerular filtration rate En (1,3), Ev (2,5,7)Higher levels of plasma creatinine with no apparent kidney damage or renal impairmentEn (1,2), Ev (5,7)Higher urea En (1,2), Ev (5,7), glucose En (1,7), potassium Ev (7), sodium En (1), and chloride Ev (5,7)Increased plasma osmolalityEn (1,2), Ev (1, 5,7,10), Evap (5)Food/waterIncreased fat storage in body tissuesEn (1,7,7⁎,4)Energy reservation during food/water scarcityFat metabolismEn (1,2,4,7), Ev (13,14)Enhanced use of metabolic waterEn (1,2,7), Ev (1,2,8, 12,13,14), Evap (1,2)Higher plasma sodiumEn (1)Tolerance of high-salt dietEn (1,6,7⁎), Ev (1,4,8,9), Evap (5)Decreased insulin secretion without diabetesEv (1)Adaptive tolerance to dehydration and starvationa The phenotypes listed may themselves be adaptive or may be part of a correlated plastic response ([6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar,7.Rocha J.L. et al.Convergent evolution of increased urine-concentrating ability in desert mammals.Mamm. Rev. 2021; (Published online March 1, 2021. https://doi.org/10.1111/mam.12244)Crossref PubMed Scopus (2) Google Scholar,41.Ali A. et al.From desert to medicine: a review of camel genomics and therapeutic products.Front. Genet. 2019; 10: 1-20Crossref PubMed Scopus (37) Google Scholar,52.Bittner N.K.J. et al.Gene expression plasticity and desert adaptation in house mice.Evolution. 2021; (Published online January 17, 2021. https://doi.org/10.1111/evo.14172)Crossref PubMed Scopus (5) Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar,84.Scholander P.F. et al.Cold adaptation in Australian aborigines.J. Appl. Physiol. 1958; 13: 211-218Crossref PubMed Scopus (86) Google Scholar, 85.Fuller A. et al.Adaptation to heat and water shortage in large, arid-zone mammals.Physiology. 2014; 29: 159-167Crossref PubMed Scopus (48) Google Scholar, 86.Ali M.A. et al.Responses to dehydration in the one-humped camel and effects of blocking the renin-angiotensin system.PLoS ONE. 2012; 7e37299Crossref Scopus (19) Google Scholar, 87.Kordonowy L. et al.Physiological and biochemical changes associated with acute experimental dehydration in the desert adapted mouse, Peromyscus eremicus.Physiol. Rep. 2017; 5: 1-8Crossref Scopus (14) Google Scholar] for more examples and [6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar,66.Schwimmer H. Haim A. Physiological adaptations of small mammals to desert ecosystems.Integrative Zoology. 2009; 4: 357-366Crossref PubMed Scopus (35) Google Scholar,85.Fuller A. et al.Adaptation to heat and water shortage in large, arid-zone mammals.Physiology. 2014; 29: 159-167Crossref PubMed Scopus (48) Google Scholar] for detailed reviews).b Superscript codes and numbers designate: En – Endurers (1, Dromedary camel, Camelus dromedarius; 2, Arabian oryx, Oryx leucoryx; 3, Black Bedouin goat, Capra aegagrus hircus; 4, Awassi fat-tailed sheep, Ovis aries; 5, Marwari sheep; 6, Desert bighorn sheep, Ovis canadensis nelsoni; 7, domestic Bactrian camel, Camelus bactrianus; 7*, wild Bactrian camel, Camelus ferus; 8, Ethiopian Somali goat, Capra aegagrus hircus; 9, Common wallaroo, Osphranter robustus; 10, Gemsbok, Oryx gazella; 11, Arabian sand gazelle, Gazella marica); Ev – Evaders (1, Golden spiny mouse, Acomys russatus; 2, Merriam's kangaroo rat, Dipodomys merriami; 3, Tarabul's gerbil, Gerbillus tarabuli; 4, Cairo spiny mouse, Acomys cahirinus; 5, Cactus mouse, Peromyscus eremicus, 6, Spinifex hopping mouse, Notomys alexis; 7, Sonoran desert mice, Mus musculus; 8, Mongolian gerbil, Meriones unguiculatus; 9, Fawn hopping mouse, Notomys cervinus; 10, Degu, Octodon degus; 11, Agile kangaroo rat, Dipodomys agilis; 12, Desert pocket mouse, Chaetodipus penicillatus; 13, Namib dune gerbil, Gerbillurus tytonis; 14, Fat sand rat, Psammomys obesus); Evap – Evaporators (1, Fennec fox, Vulpes zerda; 2, Rueppell’s fox, Vulpes rueppellii; 3, Kit fox, Vulpes macrotis; 4, Cape hare, Lepus capensis; 5, Tammar wallaby, Macropus eugenii; 6, Springhare, Pedetes capensis); H, Aboriginal Australians, Homo sapiens. Open table in a new tab Aiming to summarize and categorize decades of classic literature on desert adaptations in species occupying distinct ecological niches, Willmer et al. [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar] proposed a classification system relative to extreme temperatures and body size. Animals that can evade extreme temperatures through behavior and are physiologically adapted to minimize water loss are called ‘evaders’. Mammalian evaders include small rodents that can avoid overheating by being nocturnal and hiding in burrows during the day, and are less reliant on evaporative cooling [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. Evaders can additionally minimize water loss by concentrating highly hyperosmotic urine with low volume [7.Rocha J.L. et al.Convergent evolution of increased urine-concentrating ability in desert mammals.Mamm. Rev. 2021; (Published online March 1, 2021. https://doi.org/10.1111/mam.12244)Crossref PubMed Scopus (2) Google Scholar] and utilize water derived from fat metabolism (metabolic water) well beyond the abilities of non-desert counterparts (Table I for examples) [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar]. Animals that cannot shelter behaviorally and instead rely on morphology and physiology to minimize water loss and withstand heat are called ‘endurers’ (Table I) [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. Endurers include large ungulates and marsupials able to store heat without increasing their body temperature [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar]. For example, Dromedary camels (Camelus dromedarius) can endure temperatures exceeding 42°C, and water losses >25% of their total body weight, which are both fatal to non-desert mammals [41.Ali A. et al.From desert to medicine: a review of camel genomics and therapeutic products.Front. Genet. 2019; 10: 1-20Crossref PubMed Scopus (37) Google Scholar]. Some endurers also have lower respiratory water losses [6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar], and can store water in their rumen, gut, or intestines for water retention [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,65.Donald J. Pannabecker T.L. Osmoregulation in desert-adapted mammals.in: Hyndman K. Pannabecker T. Sodium and Water Homeostasis. Springer, 2015: 191-211Crossref Google Scholar]. It has also been proposed that Arabian Oryx (Oryx leucocoryx ) use water derived from fat metabolism and that this may account for 24% of their overall water needs [5.Johnson R.J. et al.Metabolic and kidney diseases in the setting of climate change, water shortage, and survival factors.J. Am. Soc. Nephrol. 2016; 27: 2247-2256Crossref PubMed Scopus (49) Google Scholar]. Endurers may also have enlarged body tissues that can store localized fat (e.g., fat tails in sheep, Ovis aries, and fat-filled humps of camels) to act as energy reserves during starvation [6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar,83.Atti N. et al.Performance of the fat-tailed Barbarine sheep in its environment: adaptive capacity to alternation of underfeeding and re-feeding periods. A review.Anim. Res. 2004; 53: 165-176Crossref Scopus (70) Google Scholar]. A third category was additionally proposed for medium-sized animals that are not able to avoid extreme conditions as efficiently as evaders nor withstand heat as efficiently as endurers. These are called ‘evaporators’ and comprise lagomorphs, some marsupials, and medium-sized carnivores (Table I) [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar]. Evaporator species have enlarged body extremities (e.g., large ears) to dissipate heat by conduction and use denning and nocturnal behavior to avoid extreme heat. They also have significantly low mass-adjusted nonrenal water loss [1.Ward D. The Biology of Deserts.2nd edn. Oxford University Press, 2016Crossref Google Scholar,2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar] and can minimize renal water loss by concentrating highly hyperosmotic urine significantly above the levels seen in non-desert counterparts [2.Willmer P. et al.Environmental Physiology of Animals.2nd edn. Wiley-Blackwell, 2005Google Scholar,7.Rocha J.L. et al.Convergent evolution of increased urine-concentrating ability in desert mammals.Mamm. Rev. 2021; (Published online March 1, 2021. https://doi.org/10.1111/mam.12244)Crossref PubMed Scopus (2) Google Scholar]. However, some authors have criticized this categorization due to overlap between the categories and have proposed other frameworks to synthesize desert mammalian phenotypic adaptations (e.g., [6.Rymer T.L. et al.Resilience to droughts in mammals: a conceptual framework for estimating vulnerability of a single species.Q. Rev. Biol. 2016; 91: 133-176Crossref PubMed Scopus (24) Google Scholar]).Table IClassic Physiological Systems Allowing Mammalian Survival in DesertsaThe phenotypes listed may themselves be adaptive or may be part of a correlated plastic response ([6,7,41,52,65,84–87] for more examples and [6,65,66,85] for detailed reviews)., bSuperscript codes and numbers designate: En – Endurers (1, Dromedary camel, Camelus dromedarius; 2, Arabian oryx, Oryx leucoryx; 3, Black Bedouin goat, Capra aegagrus hircus; 4, Awassi fat-tailed sheep, Ovis aries; 5, Marwari sheep; 6, Desert bighorn sheep, Ovis canadensis nelsoni; 7, domestic Bactrian camel, Camelus bactrianus; 7*, wild Bactrian camel, Camelus ferus; 8, Ethiopian Somali goat, Capra aegagrus hircus; 9, Common wallaroo, Osphranter robustus; 10, Gemsbok, Oryx gazella; 11, Arabian sand gazelle, Gazella marica); Ev – Evaders (1, Golden spiny mouse, Acomys russatus; 2, Merriam's kangaroo rat, Dipodomys merriami; 3, Tarabul's gerbil, Gerbillus tarabuli; 4, Cairo spiny mouse, Acomys cahirinus; 5, Cactus mouse, Peromyscus eremicus, 6, Spinifex hopping mouse, Notomys alexis; 7, Sonoran desert mice, Mus musculus; 8, Mongolian gerbil, Meriones unguiculatus; 9, Fawn hopping mouse, Notomys cervinus; 10, Degu, Octodon degus; 11, Agile kangaroo rat, Dipodomys agilis; 12, Desert pocket mouse, Chaetodipus penicillatus; 13, Namib dune gerbil, Gerbillurus tytonis; 14, Fat sand rat, Psammomys obesus); Evap – Evaporators (1, Fennec fox, Vulpes zerda; 2, Rueppell’s fox, Vulpes rueppellii; 3, Kit fox, Vulpes macrotis; 4, Cape hare, Lepus capensis; 5, Tammar wallaby, Macropus eugenii; 6, Springhare, Pedetes capensis); H, Aboriginal Australians, Homo sapiens.Environmental stressorUpstream phenotypeDownstream phenotypeHeat + waterDecreased thyroxinEn (4,5), Ev (2,5), HLow metabolism or energy expendituresEn (1,2,9), Ev (1,2,8), Evap (1), HLow metabolism/energy expendituresReduced nonrenal water lossEn (1,2,10), Ev (2,8), Evap (1,2), H, low respiration rateEn (1,2), Ev (6), Evap (1), torporEv (1,5)ColdNonshivering thermogenesisEv (1,8)Reduced shiveringHHeatReduced aldosteroneEn (1)Low cardiac rate and low blood pressureEn (1)WaterIncreased vasopressinEn (1,8), Ev (1,6,10); Increased aldosteroneEn (1,3,4), Ev (3,4)Increased urine osmolality/Increased water reabsorption from the kidneyEn (1,2), Ev, Evap (1,4); decreased urine productionEv (1,5,7)Reduced aldosteroneEn (1)Increased urine sodium excretionEn (1)Low glomerular filtration rate En (1,3), Ev (2,5,7)Higher levels of plasma creatinine with no apparent kidney damage or renal impairmentEn (1,2), Ev (5,7)Higher urea En (1,2), Ev (5,7), glucose En (1,7), potassium Ev (7), sodium En (1), and chloride Ev (5,7)Increased plasma osmolalityEn (1,2), Ev (1, 5,7,10), Evap (5)Food/waterIncreased fat" @default.
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• W3153073103 date "2021-07-01" @default.
• W3153073103 modified "2023-10-11" @default.
• W3153073103 title "Life in Deserts: The Genetic Basis of Mammalian Desert Adaptation" @default.
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