Naked mole-rat

The naked mole-rat (Heterocephalus glaber), also known as

Naked mole-rat
the sand puppy,[4] is a burrowing rodent native to parts of East
Temporal range: 4.3–0 Ma
Africa. It is closely related to the blesmols and is the only ↓

species in the genus Heterocephalus of the family PreЄ Є O S D C P T J K PgN

Heterocephalidae.[5] The naked mole-rat and the Damaraland Early Pliocene - Recent[1]
mole-rat (Fukomys damarensis)[6][7] are the only known
eusocial mammals, the highest classification of sociality. It has
a highly unusual set of physical traits that allow it to thrive in a
harsh underground environment and is the only mammalian
thermoconformer, almost entirely ectothermic (cold-blooded)
in how it regulates body temperature.[8]

The naked mole-rat lacks pain sensitivity in its skin, and has
very low metabolic and respiratory rates. While formerly
considered to belong to the same family as other African mole- Conservation status
rats, Bathyergidae, more recent investigation places it in a
separate family, Heterocephalidae.[9][10][11] The naked mole-rat
is also remarkable for its longevity and its resistance to cancer
and oxygen deprivation. Least Concern (IUCN 3.1)[2]
Scientific classification
Kingdom: Animalia
Contents Phylum: Chordata
Description Class: Mammalia
Physiology
Order: Rodentia
Metabolism and respiration
Thermoregulation Family: Heterocephalidae
Pain insensitivity Landry, 1957; Patterson,
Resistance to cancer 2014
Longevity
Genus: Heterocephalus
Size
Rüppell, 1842[3]
Chronobiology
Species: H. glaber
Ecology and behavior
Distribution and habitat Binomial name
Roles
Heterocephalus glaber
Queen and gestation
Rüppell, 1842[3]
Workers
Dispersers
Colonies
Female mate choice
Diet
Predators
 Conservation status
References
Further reading
External links

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 Distribution of the naked mole-rat
(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.[12] About a quarter of their
musculature is used in the closing of their jaws while they dig— Play media
Various aged naked mole-rats.
about the same proportion that is utilized in the human leg. They
have little hair (hence the common name) and wrinkled pink or
yellowish skin. They lack an insulating layer in the skin.

Physiology

Metabolism and respiration

The naked mole-rat is well adapted to the limited availability of oxygen within the tunnels of its typical
habitat. It has underdeveloped lungs and its hemoglobin has a high affinity for oxygen, increasing the
efficiency of oxygen uptake.[13][14][15] It has a very low respiration and metabolic rate for an animal of
its size, about 70% that of a mouse, thus using oxygen minimally.[16] In response to long periods of
hunger, its metabolic rate can be reduced by up to 25 percent.[14]

The naked mole-rat survives for at least 5 hours in air that contains only 5% oxygen; it does not show
any significant signs of distress and continues normal activity. It can live in an atmosphere of 80% CO2
and 20% oxygen. In zero-oxygen atmosphere, it can survive 18 minutes apparently without suffering any
harm (but none survived a test of 30 minutes). During the anoxic period it loses consciousness, its heart
rate drops from about 200 to 50 beats per minute, and breathing stops apart from sporadic breathing
attempts. When deprived of oxygen, the animal uses fructose in its anaerobic glycolysis, producing lactic
acid. This pathway is not inhibited by acidosis as happens with glycolysis of glucose.[14][15] As of April
2017, it was not known how the naked mole-rat survives acidosis without tissue damage.[17]

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."[18] 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.[19] At lower temperatures, naked mole-
rats can use behavioral thermoregulation. For example, cold naked mole-rats huddle together or seek
shallow parts of the burrows that are warmed by the sun. Conversely, when they get too hot, naked mole-
rats retreat to the deeper, cooler parts of the burrows.

Pain insensitivity
The skin of naked mole-rats lacks neurotransmitters in their
cutaneous sensory fibers. As a result, the naked mole-rats feel no
pain when they are exposed to acid or capsaicin. When they are
injected with substance P, a type of neurotransmitter, 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 Captive naked mole-rats huddling
ventilated living spaces which would cause acid to build up in together
their body tissues.[20]

Naked mole-rats' substance P deficiency has also been tied to their lack of the histamine-induced itching
and scratching behavior typical of rodents.[21]

Resistance to cancer
Naked mole-rats have a high resistance to tumours, although it is likely that they are not entirely immune
to related disorders.[22] 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.[23]

In 2013, scientists reported that the reason naked mole-rats do not get cancer can be attributed to 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.[24][25][26]
The scientific report was published a month later as the cover story of the journal Nature.[27] A few
months later, the same University of Rochester research team announced that naked mole-rats have
ribosomes that produce extremely error-free proteins.[28] Because of both of these discoveries, the
journal Science named the naked mole-rat "Vertebrate of the Year" for 2013.[29]

In 2016, a report was published that recorded the first ever discovered malignancies in two naked mole-
rats, in two individuals.[30][31][32] However, both naked mole-rats were captive-born at zoos, and hence
lived in an environment with 21% atmospheric oxygen compared to their natural 2–9%, which may have
promoted tumorigenesis.[33]

The Golan Heights blind mole-rat (Spalax golani) and the Judean Mountains blind mole-rat (Spalax
judaei) are also resistant to cancer, but by a different mechanism.[34]
Longevity
The naked mole-rat is also of interest because it is extraordinarily long-lived for a rodent of its size (up to
32 years[15][35]) and holds the record for the longest living rodent.[36] The mortality rate of the species
does not increase with age, and thus does not conform to that of most mammals (as frequently defined by
the Gompertz-Makeham law of mortality).[37] Naked mole-rats are highly resistant to cancer[38] and
maintain healthy vascular function longer in their lifespan than shorter-living rats.[39] 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".[40] Their longevity has also been attributed to "protein stability".[41]
Because of their extraordinary longevity, an international effort was put into place to sequence the
genome of the naked mole-rat.[42] A draft genome was made available in 2011[43][44][45] with an
improved version released in 2014.[46] Its somatic number is 2n = 60.[5] 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.[47]

The DNA repair transcriptomes of the liver of humans, naked mole rats and mice were compared.[48] The
maximum lifespans of humans, naked mole rats, and mice are respectively c. 120, 30 and 3 years. The
longer-lived species, humans and naked mole rats, expressed DNA repair genes, including core genes in
several DNA repair pathways, at a higher level than did mice. In addition, several DNA repair pathways
in humans and naked mole rats were up-regulated compared with mice. These findings suggest that
increased DNA repair facilitates greater longevity, and also are consistent with the DNA damage theory
of aging.[49]

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 counterparts of the same skull width. 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.[50] 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.[51]

Chronobiology
The naked mole-rat's subterranean habitat imposes constraints on its circadian rhythm.[52] Living in
constant darkness, most individuals possess a free-running activity pattern and are active both day and
night, sleeping for short periods of time several times in between.[52]

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.[53]

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.[54]

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.[55][56] Only one female (the queen) and
one to three males reproduce, while the rest of the members of the colony function as workers.[57] The
queen and breeding males are able to breed at one year of age. Workers are sterile,[56] 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 appear to be reproductively suppressed, meaning the ovaries do not
fully mature, and do not have the same levels of certain hormones as the reproducing females. On the
other hand, there is little difference of hormone concentration between reproducing and non-reproducing
males. In experiments where the reproductive female was removed or died, one of the non-reproducing
females would take over and become sexually active. Non-reproducing members of the colony are
involved in cooperative care of the pups produced by 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.[51]

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
Female mole-rat
place, sometimes after a violent struggle with her competitors.
Once established, the new queen's body expands the space
between the vertebrae in her backbone to become longer and ready to bear pups.[58]

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.[59] 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.[60] 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 fecal pap 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.[61] As in certain bee species, the workers are divided along a continuum of
different worker-caste behaviors instead of discrete groups.[54] 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.[62] Workers are sterile when there
is no new reproductive role to fill.

Dispersers
Inbreeding is a common phenomenon among naked mole-rats within a colony. Prolonged inbreeding is
usually associated with lower fitness.[63] However, the discovery of a disperser role has revealed an
outbreeding mechanism in addition to inbreeding avoidance.[64] Dispersers are morphologically,
physiologically as well as behaviorally distinct from colony members and actively seek to leave their
burrow when an escape opportunity presents itself.[64] These individuals are equipped with generous fat
reserves for their journey.[64] Though they possess high levels of luteinizing hormone, dispersers are only
interested in mating with individuals from foreign colonies rather than their own colony's queen.[64] They
also show little interest in working cooperatively with colony members in their natal burrow.[64] Hence,
the disperser morph is well-prepared to promote the exchange of individuals as well as genetic material
between two otherwise separate colonies.[64]

Colonies
Colonies range in size from 20 to 300 individuals, with an average of 75.[65]

Female mate choice

Reproductively active female naked mole rats tend to associate with unfamiliar males (usually non-kin),
whereas reproductively inactive females do not discriminate.[66] The preference of reproductively active
females for unfamiliar males is interpreted as an adaptation for inbreeding avoidance. Inbreeding is
avoided because it ordinarily leads to the expression of recessive deleterious alleles.[67]

Diet
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.
A single tuber can provide a colony with a long-term source of
food—lasting for months, or even years,[54] 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.

Naked mole-rats sometimes also eat their own feces.[54] This may
A captive naked mole-rat eating
be part of their eusocial behavior and a means of sharing
hormones from the queen.[68]

Predators
Naked mole rats are primarily preyed upon by snakes—especially the Rufous beaked snake and Kenyan
sand boa—as well as various raptors. They are at their most vulnerable when constructing mounds and
ejecting soil to the surface.[69]

Conservation status
Naked mole-rats are not threatened. They are widespread and numerous in the drier regions of East
Africa.[70]

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Further reading
Ciszek, Deborah (1999). "Heterocephalus glaber, naked mole rat" (http://animaldiversity.um
mz.umich.edu/site/accounts/information/Heterocephalus_glaber.html). Animal Diversity
Web. Retrieved 22 January 2010.
"Heterocephalus glaber: Cooperation is Key" (https://web.archive.org/web/2010120617534
1/http://www.brookfieldzoo.org/pagegen/htm/fix/fg/fg_body.asp?sAnimal=Naked+mole-rat).
Brookfield Zoo. Archived from the original (http://www.brookfieldzoo.org/pagegen/htm/fix/fg/f
g_body.asp?sAnimal=Naked+mole-rat) on 6 December 2010. Retrieved 22 January 2010.
"Naked Mole Rat" (https://www.czs.org/Brookfield-ZOO/Zoo-Animals/Fragile-Desert/Naked-
Mole-Rat). Chicago Zoological Society. Retrieved 26 April 2017.
Baskauf, Steve (2003). "Naked mole rat (Heterocephalus glaber) images" (http://www.cas.v
anderbilt.edu/bioimages/animals/mammalia/naked-mole-rat.htm). Bioimages at Vanderbilt
University. Retrieved 22 January 2010.
Bryner, Jeanna (October 2006). "Naked Mole-rats Hold Clues to Human Aging" (http://www.l
ivescience.com/humanbiology/061009_aging_rats.html). LiveScience. Retrieved 22 January
2010.
Sherman PW, Jarvis JU (November 2002). "Extraordinary life spans of naked mole-rats
(Heterocephalus glaber)". Journal of Zoology. 258 (3): 307–311.
doi:10.1017/S0952836902001437 (https://doi.org/10.1017%2FS0952836902001437).
Shuster G, Sherman PW (1998). "Tool use by naked mole-rats". Animal Cognition. 1 (1):
71–74. doi:10.1007/s100710050009 (https://doi.org/10.1007%2Fs100710050009).
Choi CQ (January 2008). "Strange Creature Immune to Pain" (http://www.livescience.com/a
nimals/080128-mole-rat-pain.html). LiveScience. Retrieved 22 January 2010.
"Small Mammals (with a naked mole-rat webcam)" (http://nationalzoo.si.edu/Animals/Small
Mammals/default.cfm?cam=NMR). Smithsonian National Zoological Park. Retrieved
22 January 2010.

External links
View the hetGla2 (https://genome.ucsc.edu/cgi-bin/hgTracks?db=hetGla2) genome
assembly in the UCSC Genome Browser.

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