House Mouse

The House Mouse (Mus musculus) is one of the most numerous species of the genus Mus commonly termed a mouse. It is a small mammal and a rodent. In most parts of the world, they live in close proximity to humans. Laboratory mice belong to strains of House Mice and are some of the most important model organisms in biology and medicine; they are by far the most commonly used laboratory mammal.

Characteristics
House mice have an adult body length (nose to base of tail) of 7.5–10 cm and a tail length of 5–10 cm; the weight is typically 10–25 g. They vary from light brown to black, with 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 long; the normal gait is a run with a stride of about 4.5 cm, though they can jump up to 45 cm. The droppings are blackish, about 3 mm long, and have a strong musty smell. The voice is a high-pitched squeak.

Young males and females are not easily distinguished; females have a significantly smaller distance between their anus and genital opening. Females have 5 pairs of mammary glands and nipples; males have no nipples. When sexually mature, the most striking and obvious difference is the presence of testicles on the males. These are relatively 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 (Terszowski 2006).

Subspecies
There are three widely accepted subspecies, increasingly treated as distinct species:
 * Mus (musculus) musculus (East European House Mouse)
 * Mus (musculus) castaneus (Southeast Asian House Mouse)
 * Mus (musculus) domesticus (West European House Mouse)

An additional subspecies was described by Prager, Orrego and Sage (1998) from the Arabian Peninsula:
 * Mus musculus gentilulus

The following were previously identified as subspecies, but have since been found to belong to the subspecies above:
 * Mus musculus homourus
 * Mus musculus molossinus (the Japanese house mouse; actually a hybrid of musculus, castaneus and domesticus; Bonhomme 1989)
 * Mus musculus bactrianus (southwestern Asian House Mouse)
 * Mus musculus praetextus
 * Mus musculus wagneri

Behaviour
House mice usually walk, run or stand on all fours, but when eating, fighting or orienting themselves, they stand only on the hind legs, supported by the tail. When running, 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 that are 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 they will also accept meat and dairy products. They will drink water but require little of it, relying mainly on the moisture present in their food. They will eat their droppings to acquire nutrients produced by bacteria in their guts. House mice, like other rodents, do not vomit.

Mice are afraid of rats, which often kill and (partially) eat them. 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 (Tattersall, Smith and Nowell 1997). However in some areas (such as Australia) mice are able to co-exist with other small rodent species (Moro and Morris 2000).

Senses and communication
As primarily nocturnal animals, house mice have little or no colour vision. They have a sharp sense of hearing and can perceive ultrasound, possibly up to 100 kHz. They communicate both in the human audible range with squeaks (for long-distance warnings), and in the ultrasound range (for short-distance communication).

House mice also rely on pheromones. Most of these are produced by the preputial glands of both sexes and are excreted with urine. The tear fluid of male mice also contains pheromones (Kimoto 2005). 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. In (Achiraman 2002), ten different compounds such as alkanes, alcohols, etc. were detected in the urine. Among the ten, 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.

The mice can sense surfaces and air movements with their whiskers.

Life cycle and reproduction
Female house mice have an estrous cycle that is 4-6 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 become estrous after 72 hours.

Male house mice court females by emitting characteristic ultrasonic calls in the 30kHz - 110kHz 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 be different in different individuals and have been compared to birdsongs because of their complexity. (Holy 2005) While females have the capability to produce ultrasonic calls, they typically do not do so during mating behaviour.

Following copulation, female mice will normally develop a vaginal 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 3-14 young (average 6-8). One female can have some 5-10 litters per year, so their population can increase very quickly. Breeding occurs throughout the year (however, animals living in the wild don't reproduce in the colder months, even though they don't hibernate). The newborn are blind and furless. Fur starts to grow some three days after birth and the eyes open one to two weeks after birth. Females reach sexual maturity at about 6 weeks and males at about 8 weeks, but both can breed as early as five weeks.

House mice usually live under a year in the wild, because of 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, the record holder was a genetically engineered mouse that lived for 1819 days, nearly 5 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 4 years.

Mice and humans
House mice usually live in proximity to humans, in or around houses or fields. Originally native to Asia (probably northern India; Boursot et al. 1996), they spread to the Mediterranean Basin about 8000 BC, only spreading into the rest of Europe around 1000 BC (Cucci, Vigne and Auffrey 2005). This time lag is thought to be because the mice require agrarian human settlements above a certain size (Cucci, Vigne and Auffrey 2005). 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, Gunduz et al 2001, showed a previously unsuspected early link between Denmark and Madeira on the basis of the origin of the Madeiran mice. House mice can transmit diseases, and can damage food and food packaging. They can also cause substantial damage when feeding on grain. It is thought that house mice were the primary reason for the taming of the domestic cat. Various mousetraps have been developed to catch mice. Generally, rats are more harmful to humans than 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 B.C. version. Human domestication led to numerous strains of "fancy" or hobby mice with a variety of colours and a docile temperament. Domestic varieties of the house mouse called "feeder" mice are also used as food for some carnivorous pet reptiles, arthropods and fish. Mice bred for this purpose are genetically identical to other domestic mice, and can be kept as pets themselves.

Mice as an invasive species
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 one metre 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 a total of over 1 million bird chicks per year.

Laboratory mice
Mice are the most commonly used animal research model with hundreds of established inbred, outbred, and transgenic strains. In the United States, they are generally not regulated under the Animal Welfare Act (AWA) administered by the USDA, APHIS. However, the Public Health Service Act (PHS) as administered by the National Institutes of Health (NIH) does cover their humane treatment. Most academic research institutes seek voluntary accreditation which requires certain minimal standards of care for laboratory animals. This accreditation is a prerequisite for federal funding.

Mice are common experimental animals in biology and psychology primarily because they are mammals, and thus share a high degree of homology with humans. The mouse genome has been sequenced, and virtually all mouse genes have human homologs. They can also be manipulated in ways that would be considered unethical to do with humans. Mice are a primary mammalian model organism, as are rats. There are many additional benefits of mice in laboratory research. Mice are small, inexpensive, easily maintained, and can reproduce quickly. Several generations of mice can be observed in a relatively short period of time. Some mice can become docile if raised from birth and given sufficient human contact. However, certain strains have been known to be quite temperamental.

Most laboratory mice are hybrids of different subspecies, most commonly of Mus musculus domesticus and Mus musculus musculus. Laboratory mice come in 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.

The behavioural patterns of laboratory mice are significantly different from those of most common house mice due to years of lab breeding. These behaviours are much simpler.



Genome
Sequencing of the mouse genome was completed in late 2002. The haploid genome is about 3 billion bases long (3000 Mb distributed over 20 chromosomes) and therefore equal to the size of the human genome. 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. 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.

Mutant and transgenic strains
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.
 * Mice with unusual regenerative capacities.
 * "Waltzing" mice, which walk in a circular pattern due to a mutation adversely affecting their inner ear.
 * Immunodeficient nude mice, lacking hair and a thymus. The mice don't produce T lymphocytes and therefore don't mount cellular immune responses. They are used for research in immunology and transplantation.
 * Severe Combined Immunodeficient or SCID mice, 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 mice, with a disabled myostatin gene.

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

External links and sources

 * Comprehensive house mouse information, including pictures, by the University of Michigan Museum of Zoology
 * High-Resolution Brain Maps and Brain Atlases of Mus musculus
 * Biology of the Mouse, from the Louisiana Veterinary Medical Association
 * Biology of Laboratory Rodents by David G. Besselsen
 * Mus musculus, by the Museum of Texas Tech University
 * Nature Mouse Special 2002
 * Ensembl Mus musculus genome browser, from the Ensembl Project
 * Vega Mus musculus genome browser, includes NOD mouse sequence and annotation
 * Taxonomy entry from NCBI, with comprehensive links to database information about Mus musculus
 * Pictures, movies and applets showing the anatomy of Mus musculus, from www.digimorph.org
 * Monica Lawlor: A Home For A Mouse, Humane Innovations and Alternatives (Vol 8, 1994). Description of behaviour and senses.
 * 'Fancy Mice', includes much behavioural and physiological information
 * Some information on muricide
 * Vocalizations during copulation
 * "Songs" of male mice
 * 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
 * "Surprise organ discovered in mice; Mice are shown to have two thymus organs, not just one" March 2, 2006, Nature online

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