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Naked mole-rat

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The naked mole-rat (Heterocephalus glaber) also known as the sand puppy or desert mole rat, is a burrowing rodent native to parts of East Africa and is the only species currently classified in the genus Heterocephalus.[2] The naked mole-rat and the Damaraland mole-rat are the only known eusocial mammals.[3] It has a highly unusual set of physical traits that enable it to thrive in an otherwise harsh underground environment; it is the only mammalian thermoconformer,[4] has a lack of pain sensation in its skin, and has very low metabolic and respiratory rates. It is also remarkable for its resistance to cancer and its longevity.

Naked mole-rat
Temporal range: Early Pliocene - Recent
Scientific classification Edit this classification
Missing taxonomy template (fix): Heterocephalus glaber
Binomial name
Heterocephalus glaber
Rüppell, 1842
Distribution of the naked mole-rat

Description

Typical individuals are 8 to 10 cm (3 to 4 in) long and weigh 30 to 35 grams (1.1 to 1.2 oz). Queens are larger and may weigh well over 50 grams (1.8 oz), the largest reaching 80 grams (2.8 oz). They are well-adapted to their underground existence. Their eyes are quite small, and their visual acuity is poor. Their legs are thin and short; however, they are highly adept at moving underground and can move backward as fast as they can move forward. Their large, protruding teeth are used to dig and their lips are sealed just behind the teeth, preventing soil from filling their mouths while digging. About a quarter of their musculature is used in the closing of their jaws while they dig - about the same proportion as found in the human leg.[5] They have little hair (hence the common name) and wrinkled pink or yellowish skin. They lack an insulating layer in the skin.

Its karyotype has 2n = 60.[2]

Physiology

The naked mole-rat is well adapted for the limited availability of oxygen within the tunnels that are its habitat: its lungs are very small and its blood has a very strong affinity for oxygen, increasing the efficiency of oxygen uptake. It has a very low respiration and metabolic rate for an animal of its size, about 2/3 that of a similarly sized mouse, thus using oxygen minimally. In long periods of hunger, such as a drought, its metabolic rate can be reduced by up to 25 percent.

Thermoregulation

The naked mole-rat does not regulate its body temperature in typical mammalian fashion. They are thermoconformers rather than thermoregulators in that, unlike other mammals, body temperature tracks ambient temperatures. However, it has also been claimed that "the Naked Mole-Rat has a distinct temperature and activity rhythm that is not coupled to environmental conditions."[6] The relationship between oxygen consumption and ambient temperature switches from a typical poikilothermic pattern to a homeothermic mode when temperature is at 28 °C or higher.[7] At lower temperatures naked mole-rats can use behavioral thermoregulation. For example, cold naked mole-rats huddle together or bask in the shallow, more sun-warmed parts of the burrows. Conversely, when they get too hot, naked mole-rats retreat to the deeper, cooler parts of the burrows.

Pain insensitivity

 
Captive naked mole-rats huddling together

The skin of naked mole-rats lacks a key neurotransmitter called substance P that is responsible in mammals for sending pain signals to the central nervous system. The naked mole-rats feel no pain when they are exposed to acid or capsaicin. When they are injected with substance P, however, the pain signaling works as it does in other mammals, but only with capsaicin and not with acids. This is proposed to be an adaptation to the animal living in high levels of carbon dioxide due to poorly ventilated living spaces, which would cause acid to build up in their body tissues.[8]

Naked mole-rats' substance P deficiency has also been tied to their lack of the histamine-induced itching and scratching behavior typical of rodents.[9]

Resistance to cancer

Naked mole-rats appear to have a high resistance to tumours; cancer has never been observed in them.[10] A potential mechanism that averts cancer is an "over-crowding" gene, p16, which prevents cell division once individual cells come into contact (known as "contact inhibition"). The cells of most mammals, including naked mole-rats, undergo contact inhibition via the gene p27 which prevents cellular reproduction at a much higher cell density than p16 does. The combination of p16 and p27 in naked mole-rat cells is a double barrier to uncontrolled cell proliferation, one of the hallmarks of cancer.[11]

On June 19, 2013, scientists reported that the reason naked mole-rats do not get cancer may be because they produce an "extremely high-molecular-mass hyaluronan" (HMW-HA) (a natural sugary substance), which is over "five times larger" than that in cancer-prone humans and cancer-susceptible laboratory animals.[12][13][14] The breakthrough scientific report was published a month later as the cover story of the journal Nature.[15] A few months later, the same University of Rochester research team announced that naked mole-rats have ribosomes that produce extremely error-free proteins.[16] Because of both of these discoveries, the journal Science named the naked mole-rat "Vertebrate of the Year" for 2013.[17]

Blind mole-rats Spalax golani and Spalax judaei also appear to be immune to cancer but by a different mechanism.[18]

Longevity

The naked mole-rat is also of interest because it is extraordinarily long-lived for a rodent of its size (up to 31 years[19]) and holds the record for the longest living rodent.[20] Naked mole-rats are highly resistant to cancer[10] and maintain healthy vascular function longer in their lifespan than shorter-living rats.[21] The reason for their longevity is debated, but is thought to be related to their ability to substantially reduce their metabolism during hard times, and so prevent aging-induced damage from oxidative stress. This has been referred to as "living their life in pulses".[22] Their longevity has also been attributed to “protein stability.”[23] Because of their extraordinary longevity, an international effort was put into place to sequence the genome of the naked mole-rat.[24] A draft genome was made available in 2011[25][26][27] with an improved version released in 2014.[28] Further transcriptome sequencing revealed genes related to mitochondria and oxidation reduction processes to have high expression levels in the naked mole-rat when compared to mice, which may contribute to their longevity.[29]

Size

Reproducing females become the dominant female, usually, by founding new colonies, fighting for the dominant position, or taking over once the reproducing female dies. These reproducing females tend to have longer bodies than that of their non-reproducing counter parts of the same skull width. Interestingly enough, the measurements of females before they became reproductive and after show significant increases in body size. It is believed that this trait does not occur due to pre-existing morphological differences but to the actual attainment of the dominant female position.[30] As with the reproductive females, the reproductive males also appear to be bigger in size than their non-reproducing counterparts but not as much so as in the case of the females. These males also have visible outlines of the testes through the skin of their abdomens. Unlike the females, there are usually multiple reproducing males [31]

Ecology and behavior

Distribution and habitat

The naked mole-rat is native to the drier parts of the tropical grasslands of East Africa, predominantly southern Ethiopia, Kenya, and Somalia.[32]

Clusters averaging 75 to 80 individuals live together in complex systems of burrows in arid African deserts. The tunnel systems built by naked mole-rats can stretch up to three to five kilometres (2–3 mi) in cumulative length.[33]

Roles

The naked mole-rat is the first mammal discovered to exhibit eusociality. This eusocial structure is similar to that found in ants, termites, and some bees and wasps.[34][35] Only one female (the queen) and one to three males reproduce, while the rest of the members of the colony function as workers. The queen and breeding males are able to breed at one year of age. Workers are sterile,[35] with the smaller focusing on gathering food and maintaining the nest, while larger workers are more reactive in case of attack. The non-reproducing females have appeared to be reproductively suppressed, meaning the ovaries do not fully mature. They also do not have the same levels of certain hormones as the reproducing females. The males, on the other hand, do not show much difference in their hormone concentrations. In experiments where the reproductive female was removed or died, one of the non-reproducing females would take over and become sexually active. Along with the reproducing female and reproducing males, there are also non-reproductive individuals that make up the colony. It can be said that there are two “jobs” that these individuals perform. The groups that perform the jobs of infrequent or frequent workers are made up of both the non-reproducing male and non-reproducing female. There is also cooperative care that occurs for the pups of the reproducing female. This occurs through the workers keeping the pups from straying, foraging for food, grooming, contributing to extension of tunnels, and keeping them warm.[31]

The Damaraland mole-rat (Cryptomys damarensis) is the only other eusocial mammal currently known.

Queen and gestation

The relationships between the queen and the breeding males may last for many years; other females are temporarily sterile. Queens live from 13 to 18 years, and are extremely hostile to other females behaving like queens, or producing hormones for becoming queens. When the queen dies, another female takes her place, sometimes after a violent struggle with her competitors. Once established, the new queen stretches the space between the vertebrae in her backbone to become longer and ready to bear pups.[36]

Gestation is about 70 days. A litter typically ranges from three to twelve pups, but may be as large as twenty-eight. The average litter size is eleven.[37] In the wild, naked mole-rats usually breed once a year, if the litter survives. In captivity, they breed all year long and can produce a litter every 80 days.[38] The young are born blind and weigh about 2 grams (0.07 oz). The queen nurses them for the first month; after which the other members of the colony feed them feces until they are old enough to eat solid food.

Workers

Smaller workers focus on acquiring food and maintaining tunnels, while the larger workers are more reactive in case of attacks.[39] As in certain bee species, the workers are divided along a continuum of different worker-caste behaviors instead of discrete groups.[33] Some function primarily as tunnellers, expanding the large network of tunnels within the burrow system, and some primarily as soldiers, protecting the group from outside predators. There are two main types of worker, the "frequent workers" who frequently perform tasks such as foraging and nest building and "infrequent workers" that show role overlap with the "frequent workers" but perform at a much slower rate.[40] Workers are sterile when there is no new reproductive role to fill.

Colonies

Colonies range in size from 20 to 300 individuals, with an average of 75.[41]

Diet

 
A captive naked mole-rat eating

Naked mole-rats feed primarily on very large tubers (weighing as much as a thousand times the body weight of a typical mole-rat) that they find deep underground through their mining operations, but also eat their own feces.[33] A single tuber can provide a colony with a long-term source of food—lasting for months, or even years,[33] as they eat the inside but leave the outside, allowing the tuber to regenerate. Symbiotic bacteria in their intestines ferment the fibres, allowing otherwise indigestible cellulose to be turned into volatile fatty acids.

Conservation status

Naked mole-rats are not threatened. They are widespread and numerous in the drier regions of East Africa.[42]

References

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  3. ^ O'Riain, M.J.; Faulkes, C. G. (2008). "African mole rats: eusociality, relatedness and ecological constraints. In J. Korb and J. Heinze (eds.), Ecology of Social Evolution, 207-223".
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