Laboratory Mice and Rats
Mary Johnson (mary at labome dot com)
Synatom Research, Princeton, New Jersey, United States
DOI
http://dx.doi.org/10.13070/mm.en.2.113
Date
last modified : 2014-08-29; original version : 2012-10-05
Cite as
MATER METHODS 2012;2:113
Abstract

A summary and overview about the mice and rats used in biomedical research, based on a survey of 303 publications.

  • C57BL/6 and BALB/c are the main mouse strains.
  • Sprague-Dawley and Wistar are the main rat strains.
  • The Jackson Laboratory, Charles River Laboratories, Taconic Farms, and Harlan Laboratories are the main suppliers.
  • Major research applications are in immunology, oncology, physiology, pathology, and neuroscience.
  • Specific strains or ES cell lines can be searched at IMSR database.

Introduction

Animal models are indispensable tools in biomedical research. They have been used since the early days of scientific discovery and still contribute greatly today to our understanding of the functions of individual genes, the mechanisms of different diseases, and the effectiveness and the toxicities of various medicines and chemicals. The genomes of many model organisms have been sequenced, and many genes have been found to be conserved between model organisms and human. Table 1 and 2 list the number of the articles annotated with specific MESH headings or search parameters related to mice in PUBMED database. The results indicate researches conducted on mice or related to mice comprise a significant portion of our collective research effort and knowledge.

Labome undertook a survey of animal models from 303 peer-reviewed formal publications to provide an overview of animal models in publications (Table 3). The publications surveyed are in a random subset of about 10,000 open-access publications which Labome has reviewed for antibody applications. In addition, 70 publications in the journal Science were included. Almost all publications are from 2008 to 2011. The results indicate that mouse is the overwhelmingly preferred laboratory animal; the most widely used mouse and rat strains are C57BL/6 mice, BALB/c mice, Sprague-Dawley rats, and Wistar rats. Other strains, such as A/J mice, CD1 mice, and ICR mice, were also used. The majority of these animals are supplied by four major providers, The Jackson Laboratory,Charles River Laboratories, Taconic Farms and Harlan Laboratories. The aforementioned animal models are used for research in immunology, oncology, physiology, pathology, and increasingly, neuroscience, some of the interesting mouse models are discussed later in this article.

Albinism of laboratory rodents: The majority of laboratory rodents are albinos, due to a common mutation in tyrosinase gene in all albino laboratory rat strains [1] and in at least some of the albino mouse strains [2]. Tyrosinase is the rate-limiting enzyme in the production of melanin pigment. The prevalence of albinism among laboratory rodents is because many of the earliest established strains were albino, and also albinism is an easy selection marker in the early days.

Type200820092010201120122013TotalPercent
Mice, inbred strains 20163 21524 24340 26008 26802 16308 135145 31.12
Mice, transgenic 12592 13362 14363 15209 15242 10168 80936 18.64
Mice, knockout 7885 8558 9172 9702 9622 6442 51381 11.83
Mice, congenic 125 91 111 88 76 42 533 0.12
Mice, none of above * 25812 27505 29962 32052 31834 19104 166269 38.29
Total 66577 71040 77948 83059 83576 52064 434264 100
Table 1. Types of mice and the frequency of citations in PUBMED bibliographic database, as of February 19, 2014. * PUBMED search parameter: mice [ mesh] NOT mice, inbred strains [ mesh] NOT mice, transgenic [ mesh] NOT mice, knockout [ mesh] NOT mice, congenic [ mesh].
Mice

Mouse is the most cited animal model in these publications. Among the 303 articles, 276 cited various strains of mice (along with 21 citing rats, and one or two for rabbits, ferrets, guinea pigs, and rhesus macaque each). This is not surprising, 99% of mouse genes have human counterparts. Mouse and its brethren, rat, are relatively inexpensive to raise and to maintain. They can reproduce quickly, thus enable researchers to study the function of particular genes through several generations of offsprings during a reasonable period of time. Their physiology and genetics have been studied extensively, and can be compared to human easily. Technologies, such as transgenic methods, have been developed over many decades to study the genetics and the functions of specific genes. Mouse models of many human diseases have also been developed to advance the studies of disease pathogenesis, and to evaluate the effectiveness and toxicities of various candidate drugs.

Inbred strains, congenic, and transgenic mice with inbred background are commonly used. An inbred strain is defined as a strain that has been through sib-mating (or its equivalent) over 20 generations, thus animals from the same inbred strain are considered as genetically identical. Congenic strains are achieved through repeated backcrosses to an inbred background strain to select a unique marker over a minimum of 10 generations. The most cited strains are C57BL/6, BALB/c, CD-1, SCID, and A/J. Only CD-1 is an outbred strain.

MeSH heading strains 2008 2009 2010 2011 2012 2013 Total Percent
Yes C57BL 11888 12993 14927 16006 16731 10597 83142 43.25
No NIH 6912 7467 8533 8753 8839 5526 46030 23.94
Yes BALB/c 5823 6317 7026 7407 7502 4241 38316 19.93
Yes ICR 1107 1084 1254 1377 1348 760 6930 3.6
Yes NOD 548 561 650 779 872 613 4023 2.09
Yes C3H 637 640 641 582 552 251 3303 1.72
Yes DBA 437 429 456 412 374 196 2304 1.2
No MOLD 295 312 357 362 359 208 1893 0.98
Yes CBA 342 336 347 356 273 165 1819 0.95
Yes 129 0 7 190 520 563 310 1590 0.83
Yes HRS 95 114 127 139 125 72 672 0.35
Yes MRL lpr 63 71 69 60 69 34 366 0.19
Yes A 73 70 65 64 34 26 332 0.17
Yes NZB 45 36 56 52 47 27 263 0.14
No A/J 53 40 47 48 25 19 232 0.12
No SJL 35 39 42 43 35 22 216 0.11
Yes AKR 36 33 46 23 25 20 183 0.1
No CAST 18 23 24 28 34 18 145 0.08
No NZW 26 22 27 18 20 11 124 0.06
No KK 22 25 17 16 28 14 122 0.06
Yes CFTR 25 21 21 15 19 7 108 0.06
No SAMR1 15 10 7 7 9 9 57 0.03
No C57L 7 5 7 5 7 3 34 0.02
Yes SENCAR 10 6 6 3 2 3 30 0.02
No NZO 2 6 1 0 3 4 16 0.01
No C57BR 0 3 1 2 1 0 7 0
Total 28514 30670 34944 37077 37896 23156 192257 100
Table 2. Mouse inbred strains and their citations in PUBMED database, as of February 19, 2014.
C57BL/6 mice

C57BL/6 mice, also called "C57 black 6" or simply "Black 6", has the advantage of strain stability and easy breeding. This is also the first mouse strain whose genome was fully sequenced in 2005, second only to the human genome. The International Mouse Phenotyping Consortium IMPC [3], launched on September 29, 2011, aims to catalog the function of every mouse gene in this strain through knockout technologies. The application of C57BL/6 mice consists of three main areas. The most common one is to serve as physiological or pathological models for in vivo experiments. Secondly, they are often applied to build transgenic mice models. Lastly, C57BL/6 mice are used as a background strain for the generation of congenics with both spontaneous and induced mutations.

Laboratory Mice and Rats figure 1
Figure 1. Female C57BL/6, 22 months of age.

The common C57BL/6 substrains include C57BL/6J (maintained at The Jackson Laboratory) and C57BL/6N (established at National Institute of Health). IMPC selected embryonic stem cells from C57BL/6N [4] [5], while Mouse Genome Sequencing Consortium [6] and Allen Brain Atlas [7] used C57BL/6J. The substrains have phenotypic differences [8] [9], and some of the underlying genetic alterations have been identified [8] [10]. A nonsynonymous mutation of serine to phenylalanine (S968F) in cytoplasmic FMRP interacting protein 2 (Cyfip2), present in C57BL/6N substrain (but not present in the other widely used B57BL/6J substrain), is responsible for the lower acute and sensitized response to cocaine and methamphetamine observed in the substrain [10].

Most C57BL/6 substrains are "genetically ablated of one of the most important mitochondrial antioxidant enzyme, transhydrogenase (gene NNT), due to a natural deletion in the exons of this gene that completely prevents the expression of the protein [11] [8] ", and the mutation is responsible for the impaired glucose clearance and resting glucose levels (higher than wild-type Nnt mice such as C57BL/6NJ) [12]. C57BL/6J mice with the Nnt mutation have a normal lifespan, are not diabetic, and have the normal diet-induced obesity response (from Jackson Laboratories).

Among the articles surveyed by Labome, a significant percentage (at least 133 publications) cited C57BL/6 mouse strain. It is thought that the C57BL/6 strain will continue to be the preferred strain, due to the fact that its genome has been sequenced, and to the concerted effort of gene functional analysis by IMPC. While the advantage of using a "standard" strain in research is apparent, interesting issues have been raised about the drawbacks of such a practice, see The Trouble With Black-6.

animalstrainnumsuppliernum
mouseC57BL/6133
The Jackson Laboratory78
Charles River21
Taconic Farms12
Harlan Laboratories10
mouseBALB/C31
The Jackson Laboratory10
Charles River9
Taconic Farms5
Harlan Laboratories3
mouseCD-19
mouseSCID8
mouseA/J4
ratSprague-Dawley10
ratWistar7
Table 3. The number of publications (num) cites each of major mouse and rat strains. Each strain includes substrains, hybrid strains, and background strains. The numbers of publications citing major suppliers are indicated as well. Due to the complexity of the animal strain/stock names used in the publications, the number for each strain is the lower bound.
C57BL/6 mice from The Jackson Laboratory

Wild type, congenic, and transgenic/knockout C57BL/6 mice from The Jackson Laboratory have been used to study T cell development [13] [14], B lymphocyte development [15], inflammation [16] [17] [16] [18] [19] [20], the regulation of PepT1 by butyrate [21], the protective role of anti-Stx2 antibody [22], the effect of circadian changes of redox state on the excitability of SCN neurons [23], the influence of resveratrol on mitochondrial biogenesis in muscle [24], the trafficking mechanisms of endosomal Toll-like receptors [25], the effect of exposure to microbes on NKT cells [26], the involvement of growth factor TGF-b3 in palatal shelf adhesion [27], fibrosis and cardiomyopathy [28], heart failure [29], pulmonary embolism [30], colitis [15], PI3K gamma activation [31], platelet function [32], graft-versus-host-disease [33] [34], early pneumonia [35], diabesity [36] [37], the cardioprotective effect of kinin B1 receptor [38], skin microbiota as a protective immunity [39], IL-17 and IL-22 production [40], smoking damage [41], chronic neuropathic pain [42], adaptive immunity [43], nNOS in the varicosities [44], and microvascular endothelial barrier dysfunction [45]. Infectious diseases and defense against the pathogens are the central themes of many publications employing C57BL/6 mice: commensal bacteria [46], cerebral malaria pathology [47], Legionella pneumophila [48], Citrobacter rodentium [49], malaria [50] [50], Salmonella enterica serotype Typhimurium infection [51], invasive Salmonella typhimurium infection [52]. Narita M et al. used C57BL/6J to study the synthesis of secretory proteins [53]. B6.129(Cg)-Tg(CAG-Bgeo/GFP)21Lbe/J mice [54] and B6.129X1-Gt(ROSA)26Sortm1(EYFP)Cos/J mice [54] were used.

strainmajor featuresadvantagesmain applications
C57BL/6 inbred, black strain stability, easy breeding physiological or pathological models for in vivo experiments, background strain for transgenics and congenics
BALB/c inbred, albino, immunodeficient easy breeding, tumor-prone hybridoma and monoclonal antibody production, research models for cancer therapy and immunology.
CD-1 outbred, albino genetic variabilitypositional cloning, genotypic selection, toxicology testing (questionable)
CB17 SCID inbred, albino no T and B cells, tumor transplantationimmunodeficient animal model for testing new cancer treatments and as hosts for human immune system tissues.
Table 4. Major mouse strains and their features, applications.
C57BL/6 mice from Charles River Laboratories

Charles River Laboratories is another major supplier of C57BL/6 mice. Charles River C57BL/6 have been used to study rodent malaria model parasite Plasmodium berghei [55], fetuin-A [56], endocrine islet cell differentiation [57], cholestatic liver disease [58], xenobiotic transporters [59], antigen presentation [60], suicide recombination in B cell homeostasis [61], Bmi1 function [62], the susceptibility to SD and neurological deficits [63], the involvement of PFK1 glycosylation in cancer cell growth [64], the germline methylation marks at imprinted genes [65], the genital HSV-2 infection [66], and ATM gene expression [67]. Most recently, Li XY et al. used adult male C57BL/6 mice to investigate the role of PKM-zeta on neuropathic pain hypersensitivity in cortex [68] and Oh JS et al. collected oocytes from C57BL/6 mice [69].

C57BL/6 mice from Taconic Farms

C57BL/6 mice from Taconic were used to study acute and chronic psychomotor stimulant response [70], the role of DPP-IV in the immune system [71], the role of ENT3 in lysosome function and macrophage homeostasis [72], the involvement of rapamycin through mTORC2 disruption in insulin resistance [73], the effect of acute gastrointestinal infection on immune response to commensals [74], skin microbiota as a protective immunity [39], murine lymphoproliferative diseases [75], effector cytolytic T lymphocytes [76], mast cell function [77], and NKT lymphocyte lineage development [78]. Round JL et al. used Taconic Farms C57BL/6 mice to investigate the role of Bacteroides fragilis in establishing host-microbial symbiosis [79]. B6.129S1-Tlr3 tm1Flv/J(Tlr3-/-) and B6.129S7-Rag1 tm1Mom (Rag1 -/-) mice from Taconic Farms were used to study skin microbiota2 [80] and B6.SJL-Ptprca(CD45.1) mice were used to perform parasitic infection [74].

C57BL/6 mice from Harlan Laboratories

Harlan Laboratories C57BL/6 mice were used to study ischemia reperfusion [81], Mycobacterium tuberculosis vaccination [82], neurotrophin expression [83], IL-1 secretion [84], bluetongue virus infection [85], the regulatory mechanism of T cell differentiation [86], and persistent viral infection and chronic inflammatory diseases [87].

C57BL/6 from other providers

Other suppliers provided C57BL/6 mice that were used to study ECE-1 [88], the effect of CSE1L/CAS on cancer cells [89], and the effect of delphilin ablation on both long-term depression induction and optokinetic response adaptation [90], glucose and lipid metabolism [91]. C57BL/6 mice with experimental autoimmune uveoretinitis from Sankyo Laboratory Service Corp were used to study the spreading of various retinal protein autoantigens [92] and to investigate the role of esterase D and brain-type creatine kinase as retinal autoantigens [93] in endogenous uveitis.

Academic or national animal facilities are also suppliers. Fogg DK et al. used mice expressing eGFP at the locus of the CX3CR1 gene (Cx3cr1gfp/+) on the CD45.2 C57BL/6 background from animal care facility of the University of Rene Descartes to identify a clonogenic bone marrow progenitor [94]. Age- and sex-matched C57BL/6 mice from National Cancer Institute served as wild type controls [95]. C57BL/6.Myd88-/- mice from University of Alabama at Birmingham were used to investigate the role of all-trans retinoic acid in regulatory T cell development [19]. Yi ZF et al. performed corneal micropocket assay and histological examination in C57BL/6 mice from National Rodent Laboratory Animal Resources in order to identify a novel peptide from human apolipoprotein and study its role in angiogenesis and tumor growth [96].

BALB/c mice

BALB/c is an albino, immunodeficient inbred strain. BALB/c mice have the characteristics of easy breeding and minimal weight variations between males and females. Much noteworthy is that the mammary tumour incidence in BALB/c mice is low, but they are very sensitive to carcinogens, and can develop lung tumours, reticular neoplasms, renal tumours, and others. In addition, mineral oil injection can readily induce plasmacytomas in BALB/c strain, and this strain has been used extensively for hybridoma and monoclonal antibody production. BALB/c mice are useful for research in cancer therapy and immunology.

Laboratory Mice and Rats figure 2
Figure 2. BALB/c mouse. Courtesy of The Jackson Laboratory

Among the publications surveyed by Labome, 31 publications used BALB/c mice for hybridoma development, monoclonal antibody production, infectious disease research, among others. BALB/c mice served as a general purpose animal model.

For example, Oakley MS et al. identified brain changes in cerebral malaria in wild type BALB/c mice from Jackson Laboratory [47]. Takeda K et al. used BALB/c mice and (B6 x BALB/c) F1 mouse from Charles River Japan to study the role of death receptor 5 mediated-apoptosis in cholestatic liver disease [58], Oliveira Gde A et al. used BALB/c mice from Charles River Laboratories to demonstrated that antiplasmodial immunity in mosquitos is mediated by peroxidase/NOX5 System-dependent epithelial nitration [97], Jagger BW et al. used JAX Mice and Services female BALB/c mice to to perform mouse infection in order to show that host response to influenza A virus was modulated by a novel protein PA-X [98], and Ekiert DC et al. studied the effectiveness of antibody CR8020 against group 2 influenza viruses in BALB/c mice from Charles River Laboratories [99]. Li Y et al. used female BALB/c mice (Charles River Laboratories) to demonstrate the function of sopB [100]. Capraro GA et al. investigated the effect of SV5-P/V gene mutations on virus growth and adaptive immune responses in BALB/c mice (Fredrick Cancer Research and Development Center) [101]. Kendirgi F et al. studied prototype linear DNA vaccines against the influenza virus type A/H5N1 in BALB/c mice (Harlan) [102]. Charles River BALB/c mice were used to perform parasite infection to study the antiplasmodial immunity in mosquitos [97].

One common application of BALB/c, same as in the case of C57BL/6, is to serve as the background strain for various gene deficiency/knockout studies. Nurieva RI et al. studied the roles of IL-21, TH1, TH2 or TH17 lineages in the generation of follicular helper T cells in STAT6-deficient and STAT4-deficient BALB/c mice from The Jackson Laboratory [13]. Tawara I et al. investigated the role of Th2 cytokine secretion in acute graft-versus-host-disease in female STAT6 deficient BALB/c (H-2d) mice (The Jackson Laboratory) [33]. Ramaprakash H et al. investigated antifungal immune responses to swollen conidia Aspergillus fumigatus in BALB/c (TLR9+/+) mice [103] and Castilow EM et al. used IFN-gamma C.129S7 Ifngtm1Ts/J-deficient BALB/C mice and BALB/cAnNCr mice to study the effects of gamma interferon during a secondary RSV infection [104]. Chung YW et al. used BALB/cJ and BALB/c congenic C.C3 female TLR-4lps-/lps- mice and wild type BALB/c mice (The Jackson Laboratory) to study the effect of probiotics on the development of experimental colitis using Toll-like receptor 4 (TLR-4) mutant (lps-/lps-) mice [105].

BALB/c mice play important roles in oncological research. Arscott WT et al. studied the suppression of neuroblastoma growth performed by dipeptidyl peptidase IV BALB/c (nu/nu) nude mice [106], Yi W et al. performed subcutaneous xenografts by using Charles River Laboratories athymic nude (Nu/Nu) mice to show that FGFR-TACC fusion happened in specific GBM patients [107] and studied the role of PFK1 glycosylation in cancer cell growth using Charles River Laboratories nude mice [64], Kessler JD et al. demonstrated that Myc-dependent cancer needs SUMOylation for tumorigenesis by performing breast cancer cell line xenografts using Harlan Labs female athymic nude Foxn1-nu mice [108], and Beauvais DM et al. applied athymic BALB/c nude mice (from Taconic) for corneal angiogenesis assays [109].

CD-1 mice

While both C57BL/6 and BALB/c mice are strains inbred to establish the genetic homogeneity, CD-1 mice stand out among the most commonly used research mice as an outbred stock (inbred mice are referred as strains, whereas outbred mice are referred to as stocks). The genetic variability in outbred research animal models, instead, can serve as an advantage in the positional cloning of quantitative trait loci and phenotypic or genotypic selection of a special trait. However, the usage of CD-1 mice ( and outbred stocks in general) in research fields such as toxicology (safety and efficacy testing), aging, and oncology have been critically assessed and may not be beneficial at all [110]. The appearance of CD-1 mice, same as the BALB/c strain, is albino.

Several publications used CD-1 mice, mainly from Charles River Laboratories, among the cohort surveyed by Labome. Ray S et al. used adult female CD1 mice to demonstrate that integration beta-catenin/Lef-1/Tcf-3 signaling with ERalpha is necessary for estrogen-dependent endogenous gene regulation in uterine biology [111]. Tang N et al. investigated the morphogenesis of lung tubes in CD-1 mice [112]. Schulz C et al. studied the specific myeloid cells development in Charles River CD1 mice [113], Ye H et al. evaluated synthetic optogenetic transcription device [114]. CD1 mice served as the source of mouse embryos [57], were used to study CstF-64 splice variants [115], and to investigate vertebrate organogenesis [116]. Ko HS et al. used CD-1 mice from Jackson Laboratory to investigate LRRK2 degradation [117].

CB17 SCID mice

CB17 SCID mouse (SCID refers to severe combined immunodeficiency) is an albino strain with spontaneous SCID mutation. The mutation prevents the development and maturation of both T and B cells. However, SCID mice have normal NK cells, macrophages and granulocytes. They share the same appearance as the normal mice. Due to the SCID mutation, the success ratio of human tumor transplantation is very high (even higher than nude mice), which makes them into a valuable immunodeficient animal model for testing new cancer treatments and as hosts for human immune system tissues.

Laboratory Mice and Rats figure 3
Figure 3. SCID mouse. From Dr. Patricia Brown, NCI, NIH.

Li ZG et al. collected and implanted tissue specimens in CB17 SCID mice (Charles River Laboratories) to study the role of androgen receptor in prostate cancer bone metastase treatment [118]. Takeda K et al. studied the role of death receptor 5 mediated-apoptosis in cholestatic liver disease in CB17 SCID mice (Charles River Japan) [58]. Dubrovska A et al. used NOD CB17-Prkdc (SCID) mice (The Jackson Laboratory) to study the role for PTEN/Akt/PI3K signaling in the maintenance and viability of prostate cancer stem-like cell populations [119] and Miyagawa Y et al. performed subcutaneous tumorigenicity assays in CB17-SCID mice (Clea Japan) to study the expression of the dickkopf family in Ewing family tumor cells [120]. Gonzalez ME et al. examined the hypothesis that deregulation of EZH2 contribute to ER negative breast cancer progression in SCID mice [121].

A/J mice

A/J mouse is another common albino model, with unique characteristics such as a late onset progressive muscular dystrophy and adrenal cortical hormone induced congenital cleft palate. Also, it has a high incidence of spontaneous lung adenomas, and this tumor can easily develop in response to carcinogens.

Labome reviewed 4 publications using A/J mouse as animal models. Takeda K et al. used A/J mice (Japan SLC) to study the role of death receptor 5 mediated-apoptosis in cholestatic liver disease [58]. A/J mice from The Jackson Laboratory was used by Losick VP et al. to investigate the role of a hemidominant Naip5 allele in immunity [48], by Sanders CJ et al. to show the role of flagellin in adaptive immunity [43], and by Neunuebel MR et al. to investigate SidD modulation of Rab1 de-AMPylation during L. pneumophila infection [122].

ICR mice

ICR strain, also albino, originated in Switzerland and was selected by Dr. Hauschka to create a fertile mouse line. This outbred strain was named afterthe Institute of Cancer Research in the USA [123] [110]. ICR mice are characterized by docile nature, high productivity, rapid growth rate and low incidence of spontaneous tumor [124]. Main suppliers of ICR mice are Taconic and Japan SLC. ICR mice are a general-purpose model, used in particular in toxicology, neurobiology, oncology, infection, pharmacology, and also in product safety testing. Takahiro et al. used ICR mice model to elucidate the pathogenesis of hemorrhagic fever with renal syndrome caused by Hantaan virus [125]. The sub-acute toxicity of S-thanatin was examined in ICR mice by continually intravenous injection to evaluate the security of S-thanatin [126]. Behavior difference among strains and substrains are common. Dizocilpine maleate (MK-801) showed differential effects of on cerebrocortical neuronal injury in C57BL/6J, NSA, and ICR mice [127]. Significant performance differences between two related outbred albino strains of mice, ICR and CD1, were observed in cognitive tasks, ICR suffers from a severe visual impairment making this strain difficult to use in Morris water maze that require good visual perception, CD1 does not suffer from grossly impaired vision but, similarly to the ICR strain, CD1 mice exhibit decreased freezing in all phases of context-dependent fear conditioning [128]. In addition, ICR mice from different sources may give rise to different, even contradictory research results [129].

Other mice strains and genetically-modified mice

Other mouse strains such as 129X1/Sv mice [130], T cell-deficient nude mice [131], F344/DuCrl2Swe mice [132], OF1 mice [114], athymic nude (Nu/Nu) mice [107], C57BL/10 mouse [133], FVB mice [133] [134], GF mice on a B6 background [26], Swiss Webster (SW) mice [26], and athymic nude Foxn1-nu mice [108], have also been used.

Genetically-defined and genetically-modified mice and rats are widely used in research to investigate the function of specific genes, and to serve as experimental models for different human disease. Thousands of such strains are available, with a myriad of gene alterations, strain selection, and various applicabilities.

P2X7 knockout mice from The Jackson Laboratory were used to study the role of the P2X7 receptor in the repair of in vivo corneal epithelial debridement wounds [135], and leptin knockout (ob/ob) mice (Jackson Laboratory) were used to determine the effect of O-GlcNAcylation on platelet function [32]. Also extensively used are mice with other gene alterations, such as Cg-Prkdcscid IL2rgtm1Wjl/SzJ (NOD/SCID-IL2R-/-) mice [136], B10.BR (H2k) mice [60], B6.129(Cg)-Tg(CAG-Bgeo/GFP)21Lbe/J mice [54], B6.129S1-Tlr3 tm1Flv/J(Tlr3-/-) mice [39], B6.129S2-Il6tm1Kopf/J (Il-6-/-) mice [39], B6.129S7-Rag1 tm1Mom (Rag1 -/-) mice [39], B6.129X1-Gt(ROSA)26Sortm1(EYFP)Cos/J mice [54], B6.SJL-Ptprca(CD45.1) mice [74], Bmal 1 +/- mice [23], Icam1-/- mouse [15], Il-1r1-/- mice [39], OT-II TCR transgenic mice [74], Rag1-/- mouse [15], Rosa26LSL-YFP reporter mice [113], Ubiquitin C-CreERT2 mice [73], IL-6-/- mice [87], db/db and db/m mice [137], TLR2-/- mouse [43], R6/2 Huntington mice [138], and OT-2 TCR transgenic, B6 Ly5.2, and 129/SvEv IFNAR1-deficient mice [139].

Laboratory Mice and Rats figure 4
Figure 4. Lepr(db-3J) and normal mouse. Lepr(db-3J) mice are "obese, hyperphagic, cold intolerant, insulin resistant, and infertile". Courtesy of The Jackson Laboratory
Rats

Another rodent animal, rat, is the second most cited animal model. Compared to mice, rats are bigger, more fierce, and more resistant against various ailments. Sprague-Dawley rat and Wistar rat are two most frequently used rat strains. Similar to the common mouse strains, these two rat strains are albino. On the other hand, both rat strains are outbred strains (most commonly used mouse strains are inbred strains). Laboratory rats are from the species Rattus norvegicus, while the laboratory mouse strains come from several different species.

Laboratory Mice and Rats figure 5
Figure 5. Sprague-Dawley rat. Courtesy of Harlan Laboratories
Sprague-Dawley rat

Sprague-Dawley rat is a hybrid albino strain with long narrow heads. It has high reproduction rate and low incidence of spontaneous tumor. Its calm temperament and easy handling are welcome features to busy scientists and animal lab technicians. Labome surveyed 8 publications using Sprague-Dawley rats as animal models, most of them were from Charles River.

In neurobiological research, Ewert TJ et al. studied the effect of post-infarct cardiac sympathetic hyperactivity on galanin expression in adult Sprague-Dawley rats [140], Surgucheva I et al. prepared rat retinal cells from Sprague-Dawley rats to evaluate gamma-synuclein as retinal ganglion cell marker [141], Zhou HL et al. investigated glial cell line-derived neurotrophic factor expression in the rostral and caudal stumps of the transected adult Sprague-Dawley spinal cords [142], and Schafe GE et al. investigated long-term potentiation in adult male naive Sprague-Dawley rats in vivo [143], Dinieri JA et al. conducted viral gene transfer studies in male Sprague-Dawley rats to research the altered sensitivity of rewarding and aversive drugs in mice with inducible disruption of CREB function within the nucleus accumbens [144], and Chen J et al. implanted male Sprague-Dawley rats with unilateral indwelling dorsomedial hypothalamus cannulae to study the role of DMH CCK in food intake [145].

Pathological investigation is another area that Sprague-Dawley rats have been utilized. Reungjui S et al. placed male Sprague-Dawley rat on different K+ diets to investigate whether hypokalemic nephropathy involves impaired renal angiogenesis [146]. Buchholz K et al. infected Sprague-Dawley rats with Plasmodium berghei to study the cellular role of Plrx in its life cycle [55]. Ben-Ami Bartal I et al. used Charles River Sprague-Dawley rats to perform behaviour experiments and found that rats have empathy and pro-social behavior [147].

Wistar rat

Wistar rat is another hybrid albino strain. It has the distinct honor of being the first rat strain developed to serve as a model animal. Sprague-Dawley rat strain was derived from this strain.

Laboratory Mice and Rats figure 6
Figure 6. Lobund-Wistar rat. From NCI, NIH.

Labome surveyed 7 publications employing Wistar rats. Woo S et al. used male Wistar rats to develop a mechanistic model for chemotherapy-induced anemia [148]. Siniscalco D et al. researched the role of caspase-7 in pain-associated apoptotic process in male Wistar rats [149]. Aprigliano I et al. obtained hepatic stellate cells from Wistar rats to investigate the effect of atorvastatin on apoptosis [150]. Maddahi A et al. induced ischemic brain injuries in male Wistar-Hanover rats (Mollegaard Breeding Centre) to study the role of the MEK/ERK pathway in receptor expression following ischemic brain injury [151]. Unkrüer B et al. induced status epilepticus in female Wistar Unilever rats to clarify the cellular distribution of YB-1 protein in adult brains of rats, macaques, and humans [152]. Matrone C et al. obtained embryos from pregnant Wistar rats to study the role of NGF or BDNF signaling in neuronal apoptotic death [153]. Higashida K et al. used Charles Rivers male Wistar rats to perform animal studies in order to investigate the influence of resveratrol on mitochondrial biogenesis in muscle [24].

Animal Suppliers
The Jackson Laboratory

The Jackson Laboratory, founded by Clarence Cook Little, who produced the inbred mouse strain C57BL, among others, is the most cited experimental murine animal supplier. More than 5,000 strains of mice are available from The Jackson Laboratory. It also maintains an integrated mouse information resource. In 2011, it distributed 3.0 million mice to "approximately 20,000 investigators (or laboratories) in more than 900 institutions, in at least 50 countries".

Laboratory Mice and Rats figure 7
Figure 7. The Jackson Laboratory

Over 157 publications surveyed by Labome cited The Jackson Laboratory as the source. It is the main source for the common strains like C57BL, BALB/c mouse strains, and rat strains, and less common strains like DBA/2J [20], C3H.SW (H-2b) [34], NOD/ShiLtJ [154], AKR/J (AKR) [42], FvB/NJ [48] [134], C3H/HeJ (C3H), C58/J (C58) and CBA/J [42], 129 X1/SvJ [99], and MRL/MpJ [155].

Laboratory Mice and Rats figure 8
Figure 8. DBA/2J mouse. Courtesy of The Jackson Laboratory

In addition, many of congenic and transgenic mouse strains also came from The Jackson Laboratory, such as CD8-KO and PFP-KO [47], ASKO [156], B6.129(Cg)-Tg(CAG-Bgeo/GFP)21Lbe/J mice [54], B6.129S2-Il6tm1Kopf/J (Il-6-/-) mice [39], B6.129X1-Gt(ROSA)26Sortm1(EYFP)Cos/J mice [54], Bmal 1 +/- mice [23], Icam1-/- mouse [15], Rag1-/- mouse [15], Rosa26LSL-YFP reporter mice [113], Ubiquitin C-CreERT2 mice [73], beta-cateninf/f, CMV-Cre, Rosa26R, Flk1LacZ/+ and TOPGAL [157], RAG-/- [50] [79], IFN-/- [50], B6.129P2-Tcrdtm1Mom/J [51], p53 heterozygous [62], C3H/HeJ-Hmx1mpe/J [158], GM-CSF/IL-3/IL-5 receptor deficient [159], TLR3KO [154], OVA323-339 TCR transgenic [160], N-Tg (Thy1-cre) 1Vln/J and Gt (ROSA) 26Sortm1 (eYFP-Cos) [161], DO11.10 TCR transgenic [162], Tlr2-/- [79], albumin-CRE [163], SIRT6-/+ heterozygous [164], dystrophin deficient [165], and Npc1 heterozygous [165],

Charles River Laboratories

Charles River has been providing research animals for more than 60 years. Among the publications Labome surveyed, 62 publications cited Charles River as the source. It is the major supplier for common strains like C57BL, BALB/c, and other strains like B6C3F1 [166], DBA/2 [58], TNF-alphaR1KO [167], B10.BR (H2k) mice [60], Nude mice [168] and NMRI [55]. In addition, it provided most CD-1 mice and rat strains among the publications surveyed.

Taconic Farms

Taconic Farms has been providing genetically-defined rats and mice for over 50 years. In the publications surveyed by Labome, 28 publications cited Taconic as the source. In addition to the common strains like C57BL, Taconic provided DPP-IV-/- mice [71], RAG-2-/- mice [40], [162], RAG1-/- mice [66], Egr1-/- mice [77], [78], GF mice on a B6 background [26], Swiss Webster (SW) mice [26], B6.129S1-Tlr3 tm1Flv/J(Tlr3-/-) mice [39], B6.129S7-Rag1 tm1Mom (Rag1 -/-) mice [39], Il-1r1-/- mice [39], B6.SJL-Ptprca(CD45.1) mice [74], OT-II TCR transgenic mice [74], NCR nude mice [169], and TNFalpha transgenic mice [170].

Harlan Laboratories

Twenty-four publications cited Harlan Laboratories as the supplier for their animals. The famed breeding facility, Sprague-Dawley farm in Madison, Wisconsin, where the namesake rat strain, Sprague Dawley rat, was initially bred, is now part of Harlan. Harlan supplied Hsd nude mice [169], Dark Agouti rat [171], C57BL mice [86] [133], FVB mouse [133], athymic nude Foxn1-nu mouse [108], WKT rat and LEW rat [172], and SJL/J mice [87].

Other suppliers

Japan SLC provided ICR mice [173], B6 mice [174], and B10.D2 mice [159].

Other suppliers include government agencies such as National Cancer Institute [96], organizations like Texas Institute of Genomic Medicine [175], and commercial suppliers geneOway [176], JAX Mice and Services [98], Lexicon Pharmaceuticals Inc and MSD Pharmaceutical [177].

Web resources - Where to search for a specific mouse strain?
International Mouse Strain Resource (IMSR)

IMSR is a collaboration of a dozen international mouse repositories, containing information for 26,988 strains and 210,032 EC cell lines as of February, 2014. For an example, here are 158 strains for mouse p53 gene. The website is supported by an NIH grant.

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