House mouse
Conservation status
Least Concern (IUCN 3.1)[1]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Rodentia
Family:	Muridae
Subfamily:	Murinae
Genus:	Mus
Subgenus:	Mus
Species:	M. musculus
Binomial name
Mus musculus Linnaeus, 1758
Subspecies
Mus musculus bactrianus Mus musculus castaneus Mus musculus domesticus Mus musculus gentilulus Mus musculus musculus
House Mouse range

The house mouse (Mus musculus) is a small rodent, a mouse, one of the most numerous species of the genus Mus.

As a wild animal, the house mouse mainly lives associated with humans, causing damage to crops and stored food.

The house mouse has been domesticated as the pet or fancy mouse, and as the laboratory mouse which is one of the most important model organisms in biology and medicine. It is by far the most commonly used genetically altered laboratory mammal.^[2]

Characteristics[link]

House mice have an adult body length (nose to base of tail) of 7.5–10 cm (3.0–3.9 in) and a tail length of 5–10 cm (2.0–3.9 in). The weight is typically 10–25 g (0.4–0.9 oz). They vary in colour from white to grey to light brown to black. They have short hair and a light belly. The ears and tail have little hair. The hind feet are short compared to Apodemus mice, only 15–19 mm (0.59–0.75 in) long; the normal gait is a run with a stride of about 4.5 cm (1.8 in), though they can jump vertically up to 45 cm (18 in).^[3] The voice is a high-pitched squeak.^[4][5] House mice thrive under a variety of conditions: they are found in and around homes and commercial structures, as well as in open fields and agricultural lands.

Reconstructed image of a house mouse skull

Young males and females are not easily distinguished: like people, females have a significantly smaller distance between their anus and genital opening. Females have five pairs of mammary glands and nipples; males have no nipples.^[6] When sexually mature, the most striking and obvious difference is the presence of testicles on the males. These are large compared to the rest of the body and can be retracted into the body. In addition to the regular pea-size thymus organ in the chest, house mice have a second functional pinhead-size thymus organ in the neck next to the trachea.^[7]

Subspecies[link]

The three widely accepted subspecies are increasingly treated as distinct species:^[8][9]

• Mus musculus castaneus (southern and southeastern Asia)
• Mus musculus domesticus (western Europe, southwestern Asia, Americas, Africa, and Oceania)
• Mus musculus musculus (eastern Europe and northern Asia)

Two additional subspecies have been recognized more recently:^[9]

• Mus musculus bactrianus (central Asia)
• Mus musculus gentilulus (Arabian Peninsula; Madagascar)^[10]

Many more names have been given to house mice, but are now regarded as synonyms of other subspecies. Some populations are hybrids of different subspecies, including the Japanese house mouse (M. m. molossinus).^[9][11]

Behaviour[link]

Eating

Video of a Japanese pet mouse eating at Ueno Zoo

House mice usually run, walk, or stand on all fours, but when eating, fighting, or orienting themselves, they stand only on the hind legs, supported by the tail. When they run, the horizontal tail serves for balance; the end stands up vertically, unless the mouse is frightened. Mice are good jumpers, climbers, and swimmers.

Mice are mostly active during dusk or night; they do not like bright lights. They live in a wide variety of hidden places near food sources, and construct nests from various soft materials. Mice are territorial, and one dominant male usually lives together with several females and young. Dominant males respect each other's territory and normally enter another's territory only if it is vacant. If two or more males are held together in a cage, they will often turn aggressive unless they have been raised together from birth.

House mice primarily feed on plant matter, but are omnivorous. They will eat their droppings to acquire nutrients produced by bacteria in their intestines. House mice, like most other rodents, do not vomit.

Mice are afraid of rats, which often kill and (partially) eat mice. This rat behaviour is known as muricide. Despite this behaviour, free-living populations of rats and mice do exist together in forest areas in New Zealand, North America and elsewhere. House mice are generally poor competitors and in most areas cannot survive away from human settlements in areas where other small mammals, such as wood mice, are present.^[12] However, in some areas (such as Australia), mice are able to coexist with other small rodent species.^[13]

Senses and communication[link]

Vision[link]

As primarily nocturnal animals, house mice have little or no colour vision. Their visual apparatus is basically similar to humans, but differs markedly in at least one respect. The ventral area of the mouse retina has a much greater density of ultraviolet-sensitive cones than other areas of the retina, although the biological significance of this structure is unknown.^[14][15][16]

Hearing[link]

Mice can hear over a broad spectrum of frequencies, from 80 Hz to 100 kHz (i.e. into the ultrasound range), but are most sensitive in the 15–20 kHz range and around 50 kHz. They communicate both in the human audible range with squeaks (for long-distance warnings), and in the ultrasound range (for short-distance communication).^{[citation needed]}

Olfaction[link]

House mice also rely on pheromones for social communication, some of which are produced by the preputial glands of both sexes. The tear fluid and urine of male mice also contains pheromones, such as major urinary proteins.^[17][18] Mice detect pheromones mainly with the vomeronasal organ (Jacobson's organ), located at the bottom of the nose.

The urine of house mice, especially that of males, has a characteristic strong odor. At least 10 different compounds, such as alkanes, alcohols, etc., are detectable in the urine. Among them, five compounds are specific to males, namely 3-cyclohexene-1-methanol, aminotriazole (3-amino-s-triazole), 4-ethyl phenol, 3-ethyl-2,7-dimethyl octane and 1-iodoundecane.^[19]

Odours from adult males or from pregnant or lactating females can speed up or retard sexual maturation in juvenile females and synchronise reproductive cycles in mature females (i.e. the Whitten effect). Odours of unfamiliar male mice may terminate pregnancies, i.e. the Bruce effect.

Touch[link]

Mice can sense surfaces and air movements with their whiskers which are also used during thigmotaxis. If mice are blind from birth, super-normal growth of the vibrissae occurs presumably as a compensatory response,^[20] or if the vibrissae are absent, the use of vision is intensified.^[21]

Life cycle and reproduction[link]

A seven-day-old mouse suckling on an anaesthetized mother

A baby mouse, four days old

A two-week-old, just about to open its eyes

Female house mice have an estrous cycle about four to six days long, with estrus itself lasting less than a day. If several females are held together under crowded conditions, they will often not have an estrus at all. If they are then exposed to male urine, they will come into estrus after 72 hours.

Male house mice court females by emitting characteristic ultrasonic calls in the 30 kHz–110 kHz range. The calls are most frequent during courtship when the male is sniffing and following the female; however, the calls continue after mating has begun, at which time the calls are coincident with mounting behaviour. Males can be induced to emit these calls by female pheromones. The vocalizations appear to differ between individuals and have been compared to bird songs because of their complexity.^[22] While females have the capability to produce ultrasonic calls, they typically do not do so during mating behavior.

Following copulation, female mice will normally develop a copulation plug which prevents further copulation. This plug stays in place for some 24 hours. The gestation period is about 19–21 days, and they give birth to a litter of three to 14 young (average six to eight). One female can have five to 10 litters per year, so the mice population can increase very quickly. Breeding occurs throughout the year (however, animals living in the wild do not reproduce in the colder months, even though they do not hibernate). The newborn are blind and without fur. Fur starts to grow about three days after birth, and the eyes open one to two weeks after birth. Males reach sexual maturity at about six weeks and females at about eight weeks, but both can breed as early as five weeks.

Sleep[link]

This section requires expansion.

The average sleep time of a captive House mouse is said to be 12.5 hours per day.^[23]

Life expectancy[link]

House mice usually live under a year in the wild, due to a high level of predation and exposure to harsh environments. In protected environments, however, they often live two to three years. The Methuselah Mouse Prize is a competition to breed or engineer extremely long-lived laboratory mice. As of 2005^[update], the record holder was a genetically engineered mouse that lived for 1819 days (nearly five years). Another record holder that was kept in a stimulating environment but did not receive any genetic, pharmacological, or dietary treatment lived for 1551 days (over four years).

Mice and humans[link]

House mice usually live in proximity to humans, in or around houses or fields. Originally native to Asia (probably northern India),^[24] they spread to the Mediterranean Basin about 8000 BC, only spreading into the rest of Europe around 1000 BC.^[25] This time lag is thought to be because the mice require agrarian human settlements above a certain size.^[25] They have since been spread to all parts of the globe by humans.

Many studies have been done on mouse phylogenies to reconstruct early human movements. For example, one study showed a previously unsuspected early link between Denmark and Madeira on the basis of the origin of Madeiran mice.^[26]

An individually ventilated and sealed cage for laboratory mice

House mice can transmit diseases, and can damage food and food packaging. Some of the diseases the house mouse carries can be deadly: for example, leptospirosis, murine typhus, rickettsialpox, tularemia, lymphocytic choriomeningitis ^[27] and potentially bubonic plague.^{[citation needed]} House mice can also cause substantial damage when feeding on grain. House mice were thought to be the primary reason for the taming of the domestic cat. Various mousetraps have been developed to catch mice.

The first written reference to mice kept as pets occurs in the Erya, the oldest extant Chinese dictionary, from a mention in an 1100 BC version.^[28] Human domestication led to numerous strains of "fancy" or hobby mice with a variety of colours and a docile temperament.^[29] Domestic varieties of the house mouse, called "feeder" mice, are also used as food for some carnivorous pet reptiles, birds, arthropods, and fish. Mice bred for this purpose are genetically identical to other domestic mice, and they can be kept as pets themselves.^[29]

Mice as an invasive species[link]

Gough Island in the South Atlantic is used by 20 species of seabird for breeding, including almost all of the world's Tristan albatross (Diomedea dabbenena) and Atlantic petrel (Pterodroma incerta). Until house mice arrived on the island in the 19th century with seamen, the birds did not have any mammalian predators. The mice have since grown unusually large and have learned to attack albatross chicks, which can be nearly 1 m tall, but are largely immobile, by working in groups and gnawing on them until they bleed to death. The estimated 700,000 mice on the island kill over one million bird chicks per year.^[30]

Laboratory mice[link]

Laboratory mouse (note the ear tag)

Mice are the most commonly used mammalian research model with hundreds of established inbred, outbred, and transgenic strains. They are common experimental animals in biology and psychology, primarily because they are mammals, are relatively easy to maintain and handle, reproduce quickly, and share a high degree of homology with humans. The mouse genome has been sequenced, and many mouse genes have human homologues. In addition to being small, relatively inexpensive, and easily maintained, there are further benefits to the use of mice in laboratory research. Because mice can reproduce quickly, several generations of mice can be observed in a relatively short period of time.^[31]

Most laboratory mice are hybrids of different subspecies, most commonly of Mus musculus domesticus and Mus musculus musculus. Laboratory mice can have a variety of coat colours, including agouti, black and albino. Many (but not all) laboratory strains are inbred, so as to make them genetically almost identical. The different strains are identified with specific letter-digit combinations; for example C57BL/6 and BALB/c. The first such inbred strains were produced by Clarence Cook Little in 1909. Little was influential in promoting the mouse as a laboratory organism.^[32]

Albino laboratory mice

Legislation of use in science[link]

In the United States, laboratory mice are not regulated under the Animal Welfare Act administered by the USDA APHIS. However, the Public Health Service Act (PHS) as administered by the National Institutes of Health does offer a standard for their care and use. Compliance with PHS is required to receive federal funding. PHS policy is administered by the Office of Laboratory Animal Welfare. Many academic research institutes seek accreditation voluntarily, often through Association for Assessment and Accreditation of Laboratory Animal Care, which maintains the standards of care found within The Guide for the Care and Use of Laboratory Animals and the PHS policy. This accreditation is voluntary, not a prerequisite, for federal funding.^[33] In the UK, as with all other vertebrates and some invertebrates, any scientific procedure which is likely to cause lasting distress or suffering is regulated by the Home Office under the Animals (Scientific Procedures) Act 1986.

Genome[link]

Mice genetically engineered to express green fluorescent protein under UV light

Sequencing of the mouse genome was completed in late 2002. The haploid genome is about three billion bases long (3000 Mb distributed over 20 chromosomes), therefore equal to the size of the human genome.^[34] Estimating the number of genes contained in the mouse genome is difficult, in part because the definition of a gene is still being debated and extended. The current estimated gene count is 23,786.^[35] This estimate takes into account knowledge of molecular biology, as well as comparative genomic data. For comparison, humans are estimated to have 23,686 genes.^[36]

T-haplotype[link]

The t-haplotype is a selfish element that works to disable the function of the wild house mouse, Mus musculus, sperm to ensure the fertilization of the female egg with their own sperm. This gamete killer designed and structured to suppress recombination of genes, is a single unit, of linked genes, located near the centromere of chromosome 17 and is approximately 30-40 Mb long.^[37] One part of the t-haplotype is the responder-insensitive allele Tcr. Tcr gives protection from the distorting drivers, since it shows haploid-specific expression, which means only sperm that carry the haploid are rescued from being killed.^[38] While advantageous, the frequency of this selfish element is reported as low.^[39]

The low t–haplotype frequency in the wild house mouse population is a paradox. Although +/t male mice carry equal ratios of both gamete types (+ and t) and the wild type chromosome becomes functionally inactivated as well as 90% of the offspring inherit the t chromosomes, wild house mouse populations have remained polymorphic.^[39] It would be hypothesized that the high transmission distortion ratio of t-haplotypes would become fixed in a natural population, if 90% of the offspring were inheriting the t chromosome, but due to several factors, this is not the case. Investigations of different subspecies mice populations in different locations have found there were low t frequencies in enclosure populations, as well as in different subspecies. Huang et al. (2001),^[40] in Taiwan, observed a low frequency in the subspecies, Mus castaneus, which has also been observed by both Ardlie and Silver (1998)^[39] and Carroll et al. (2004).^[41] Based on these findings, the general mechanisms of these low t frequencies in mouse populations are similar across subspecies and geographical location, making the unraveling of this paradox beneficial to not only the Mus domesticus species of mouse, but also for other species of mouse, such as Mus castaneus.

Explanations for the low frequency of t-haplotypes include factors such as population size, inbreeding, heterozygosity, and polyandry in the wild house mouse population.

Mutant and transgenic strains[link]

A knockout mouse (left), a model of obesity, compared with a normal mouse

Various mutant strains of mice have been created by a number of methods:

• Mice resulting from ordinary breeding
  • NOD mice, which develop diabetes mellitus type 1.
  • MRL mice with unusual regenerative capacities^[42]
  • "Waltzing" mice, which walk in a circular pattern due to a mutation adversely affecting their inner ears
  • Immunodeficient nude mice, lacking hair and a thymus: The mice do not produce T lymphocytes, therefore do not mount cellular immune responses. They are used for research in immunology and transplantation.
  • Severe combined immunodeficient, with an almost completely defective immune system
• Transgenic mice, with foreign genes inserted into their genome
  • Abnormally large mice, with an inserted rat growth hormone gene
  • Oncomice, with an activated oncogene, so as to significantly increase the incidence of cancer
  • Doogie mice, with enhanced NMDA receptor function, resulting in improved memory and learning
• Knockout mice, where a specific gene was made inoperable by a technique known as gene knockout: The purpose is to study the function of the gene's product or to simulate a human disease.
  • Fat mice, prone to obesity due to a carboxypeptidase E deficiency
  • Strong muscular mice, with a disabled myostatin gene, nicknamed "mighty mice."

Since 1998, it has been possible to clone mice from cells derived from adult animals.

Injection procedures[link]

Routes of administration of injections in laboratory mice are mainly subcutaneous, intraperitoneal and intravenous. Intramuscular administration is not recommended due to small muscle mass.^[43] Intracerebral administration is also possible. Each procedure has recommended injection site, approximate needle gauge and recommended maximal injected volume at a single time at one site, as given in table:

To facilitate intravenous injection into the tail, laboratory mice can be carefully warmed under heat lamps to vasodilate the vessels.^[43]

Anesthesia[link]

A common regimen for general anesthesia for the house mouse is ketamine (in the dose of 100 mg per kg body weight) plus xylazine (in the dose of 5–10 mg per kg), injected by the intraperitoneal route.^[45] It has a duration of effect of about 30 minutes.^[45]

Notes[link]

References[link]

External links[link]

Wikimedia Commons has media related to: Mus musculus

Wikispecies has information related to: Mus musculus

Taxonomy

• House mouse at the Encyclopedia of Life
• http://www.findmice.org/

Genetics

• House mouse at National Center for Biotechnology Information (NCBI)
• Ensembl Mus musculus genome browser, from the Ensembl Project
• Vega Mus musculus genome browser, includes NOD mouse sequence and annotation

Media

• Pictures, movies and applets showing the anatomy of Mus musculus, from www.digimorph.org
• Michael Purdy: "Researchers add mice to list of creatures that sing in the presence of mates"-Study of male mouse "song" with mouse song recording (MP3), by Washington University Medical School
• Arkive Photographs.Short text.
• High-Resolution Brain Maps and Brain Atlases of Mus musculus

Further reading

• Fact Sheet on House Mice Includes brief overview of habits, habitat, threats and prevention tips
• Biology of the Mouse, from the Louisiana Veterinary Medical Association
• Nature Mouse Special 2002
• Biology of Laboratory Rodents by David G. Besselsen
• Comprehensive house mouse information, including pictures, by the University of Michigan Museum of Zoology
• 'Fancy Mice', includes much behavioral and physiological information
• Some information on muricide
• Vocalizations during copulation