10

The Laboratory Rat,

Rattus norvegicus
JOY E. PALM

The common laboratory rat, Rattus norvegicus, has had a colorful

history in its progression from feral origins to the research laboratory. Ap-
parently indigenous to the steppes of Asia, the species is thought to have
migrated down the Norwegian peninsula and arrived in England via ship
in the early 18th century (Donaldson, 1924; Castle, 1947; Richter, 1954).
From there, R. norvegicus spread rapidly throughout Europe, gradually
replacing and pushing southward its plague-carrying predecessor, the
smaller, common black rat, Rattus rattus or Rattus alexandrine. The lat-
ter, which had apparently arrived in Europe more directly from India via
the Mideast between 400-1100 A.D. (Zinsser, 1935), prefers a warmer
climate and is still the most commonly encountered rat in southern
latitudes. The Norway rat arrived, again as unwelcome ship cargo, in
North America in the late 18th century. As in Europe, it gradually dis-
placed R. rattus, which was forced southward except for the few isolated
northern pockets which still exist (Castle, 1947).
The actual albino ancestors of the laboratory rat probably were
collected by fanciers in England in the 19th century to be kept and sold as
pets. According to Richter (1954), a major source of albino rats was from
the "sport" of ratbaiting, then a popular pastime in England. Albinos,
which apparently occur fairly regularly in the wild (Castle, 1947), were

JOY E. PALM-Wistar Institute, Philadelphia, Pennsylvania. Deceased, February to, 1975.

243
R. C. King (ed.), Handbook of Genetics
© Plenum Press, New York 1975
244 o. Mammals

not used in the games, and they were culled from the wild rats caught and
maintained for the sport to become collectors' items. An early report of the
use of albino rats in research was made in 1856 by the famous French
physiologist, Philipeaux, who used the species for studies on adre-
nalectomy (Richter, 1954).
Although domesticated laboratory rat strains in use today were
derived originally from wild rats of both European and North American
sources, most can probably be traced back to the colony of English albinos
brought to the Wistar Institute by Donaldson in 1906 (Donaldson, 1924).
The commercial colony of the Institute functioned until 1950, and during
that time the distribution of the common laboratory rat was world-wide
(Farris and Giffith, 1949). Consequently, there is some degree of common
ancestry with respect to the gene pool of many of the inbred rat strains
available today, regardless of their current residence. This is well
illustrated by the fact that in many instances, despite more than a half
century of separate, random breeding before the lines were fixed by in-
breeding, there is a paucity of alleles at the major histocompatibility locus
among Wistar-descended strains (Palm, 1971 b; Palm and Wilson, 1973).
In a recent compilation of inbred strains currently available (Festing and
Staats, 1973), over 50 percent of the listed strains are descendants of
Wistar albinos. In addition, numerous colonies of noninbred "Wistar"
rats, or commercial lines with partial Wistar ancestry such as Sprague-
Dawley and Long-Evans, have been a major source of rats for research.
The ancestry of the Long-Evans line, of which several inbred strains and
random-bred stocks still exist, also includes a few captive Norway rats.
One commonly used strain, the so-called Brown Norway (BN) rat
however, descends directly from captive Norways. The chocolate coat-
color mutation appeared in a colony of wild rats (captured by Helen Dean
King in 1917) and the brown animals were kept separate from 1919 until
inbred by Billingham and Silvers (1959). The same captive Norway popu-
lation was the source of much of the genetic material that allowed the
progress made by King and Castle in analyzing rat genetics (King, 1939).
Breeding experiments with albinos occurred as early as 1880, when
an English investigator named Crampe reported attempts to determine the
basis of inheritance of coat-color phenotypes, of which three were known:
albino, black, and hooding (Castle, 1947). Genetic studies, initiated after
the formal acceptance of Mendelian principles, involved the same three
coat-color phenotypes. In 1906, after analyzing Crampe's records, Don-
caster supported the postulated three "units" of heredity as the genes for
albino (c), non-agouti (a), and hooded (h) (Doncaster, 1906). A com-
bination of these three genes apparently occurred in all of the foundation
stocks brought to this country from England (Castle, 1947), and are found
together in many of the inbred strains used today.
70. The Laboratory Rat, Rattus norvegicus 245

The ensuing genetic studies were generally unsuccessful in es-

tablishing chromosomal linkage until sufficient numbers of additional
mutant genes were available; the first example of linkage was reported by
Castle and Wright in 191 S. These investigators demonstrated that two
genes for yellow coat (p, pink-eyed yellow and r, red-eyed yellow) were on
the same chromosome, and shortly thereafter the relationship of both to
the albino gene was recognized, thus establishing the first chromosomal
linkage group (LG-1) in the rat. Information on linkage grew rapidly
during the next 20-25 years, but the list of 23 mutants and 4 linkage
groups summarized in 1947 (Castle, 1947) remained fairly static until
recently. Interest in immunogenetics and biochemical genetics has now
resulted in the detection of at least 20 new genetic loci and permitted the
designation of three new linkage groups (Gasser et at., 19736; Palm,
1974). In addition, a number of mutant genes for which genetic studies
remain to be done are listed in the footnote to Table 1.
Although the early development of at least six inbred strains (Curtis et
at., 1933) provided optimum genetic material for assessing linkage of some
genetic loci (notably coat-color genes) (Curtis and Dunning, 1937, 1940),
the richest source of deviant genes was the heterogenous, noninbred
colonies of domesticated and captive wild rats. The latter, bred either inter
se or outcrossed to albinos (King, 1932), yielded 12 of the 23 mutations
listed by Castle in a 1947 summary [see also, Griineberg (1947), Farris
(1950), and Farris and Griffith (1949)].
The use of such heterogeneous stocks for studying linkage resulted in
discrepancies between independent investigations. The heroic task of
compiling most, if not all, published genetic studies of the Norway rat,
and reanalyzing the data pooled from linkage studies for each locus, has
been admirably performed by Robinson (1960, 1965, 1972), to whose
references the reader's attention is enthusiastically directed for detailed
descriptions and histories of the mutations listed in Table 1. The objective
of this report is to provide a brief historical survey of mutant rat genes
(many of which are apparently no longer available), and to list new muta-
tions not previously reviewed (Table 1). Two categories of mutations that
are currently of much interest have been listed separately-the
biochemical mutants detected as electrophoretic variants, and those
characterized as immunogenetic mutants (antigens, immunoglobulins, and
immune response genes) (Table 2). Some of those grouped in Table 2
have permitted the designation of new linkage groups (Table 3), and most
are relevant to several research disciplines, especially immunobiology,
molecular biology, and neoplasia.
The mutants listed in Tables 1 and 2 have been all subjected to
genetic analysis. Table 1 includes, when known, the linkage group of
each. The references usually are to the latest report on the mutant in
 t-:l
~
c:Tl

TABLE 1. Morphological and Histological Loci of the Norway Rata

Mutation
currently
Mutant classification Mutant description Linkage group Reference available

Behavior 'Sr, shaker 2 Castle et al. (1955) ?

w, waltzing 1 Castle (1944), Whittinghill (1944) ?
Wo, wobbly Unknown Castle et at. (1941) ?

Biochemical j, jaundice Unknown Gunn (1938), Gruneberg (1947) Yes

Coat color a, non-agouti 4 King (1939) Yes

b, brown 2 King and Castle (1935) Yes
c, albino 1 Castle (1947) Yes
Cd, ruby-eyed dilute King (1939) ?
c h , Himalayan Moutier et at. (1973d) Yes
d, dilute Unknown Curtis and Dunning (1940) ?
f, fawn 4 Castle and King (1947b) ?
Hre, restricted 7 Gumbreck et at. (1971) Yes
hi, Irish spotting
Curtis and Dunning (1937) Yes
h, hooding ;}
hn, notch 7 Castle (1951) ? 9
h r, restricted 7 Macy et al. (1972) Yes S:
p, pink-eyed yellow Castle (1919) Yes '"33
r, red-eyed yellow 1 Castle (1919) ?
s, silvering 2 Castle (1953) Yes '"0:;-
 Hypotrichosis fz, fuzzy Palm (1974) Yes ~
hr, hairless Unknown Roberts et al. (1940) ?
n, naked Unknown Castle et al. (1955) ? ;;:l
'"t'-<
.,
c-
Morphological Ca, cataract Unknown Smith and Barrentine (1943) o
fa, fatty Unknown Zucker and Zucker (1961) Yes ~
in, incisorless Greep (1941)
a
2 y
st, stub tail 3 Ratcliffe and King (1941) .,::tl
~-
Pelage variation Cu-l, curly hair 2 King (1932), Keeler and King (1942) ::tl
~
Cu-2, curly hair Unknown Gregory and Blunn (1936) Yes 2'
cw, cowlick Unknown Castle et al. (1955) ? '"
;:l
o
Sh, shaggy 2 Castle and King (1947a) ? C:!
k, kinky 3 Feldman (1935) ? ~
.: .
"'"
Physiological an, anemia 2 Smith and Bogart (1939) ?
Dx, anaphylactoid reaction Unknown Harris et al. (1963) Yes
hd, hypodactyly Unknown Moutier et al. (l973c) Yes
19, cartilage hypertrophy 1 Griineberg (1939) ?
N e, hydronephrosis Unknown Lozzio et al. (1967) Yes
re, retinal dystrophy 2 Bourne and Gruneberg (1939) Yes
Rw, warfarin resistance Greaves and Ayres (1969) Yes

a Additional eXIstmg mutants for which genetic studies remain to be performed include: mask (mk), spastic (sP), chubby (ch), crawler (cr). and diabetes insipidus
(di) (C. K. Hansen, unpublished).

t<:>
~
"'"
 TABLE 2. Electrophoretic-Variant and Immunogenetic Loci of the Norway Rat

Classification Designation Description References

t-:;j

Electrophoretic-variant Es-I Pre-albumin esterase Womack (1973) CO

"'"
loci
Es-2 Albumin -associated esterase Gasser et at. (1973b), Moutier et al. (1973a),
Womack (1973)
Es-3" Multiple organ esterases Same as Es-2
Es-4" Kidney esterase Womack (1973), Moutier (1973a)
Gl-I Plasma protein Moutier et at. (1973b)
Hb-I Hemoglobin Brdicka (1968), French et at. (1971)
Hbb Hemoglobin ,8-chain Brdicka (1968)
Svp Seminal vesicle protein Gasser (1972), Moutier et at. (l973a)

Immunogenetic loci Ag-A Natural hemagglutinin Burhoe (1947), Keeler (1949)

Ag-B" (also R- Histocom patibility Bogden and Aptekman (1960), Palm
1, RtH-1 (1971a,b), Stark et at. (1969), Palm and
Wilson (1973)
Ag-C Hemagglutinin (formerly C, D blood Owen (1962), Palm (1971a,b)
types
Ag-D Hemagglutinin Palm (1971a), Palm and Black (1971)
Ag-E Histocom patibility Michie and Anderson (1966), Palm (1971a)
Ag-F Lymphocytotoxin
Ag-G Lymphocytotoxin } C. W. DeWitt (unpublished)
Ag-Y Histocom patibility (male-limited) Gasser and Silvers (1972), Hausman and
Palm (1973a,b)
EAE-I Allergic encephalomyelitis Gasser et at. (1973a), Williams and Moore
susceptibility (1973)
IR-I Immune response'" Wurzburg (1971), Gunther et at. (1972)
~
RI-I Immunoglobulin light chain Guttman and Weisman (1971)
Ia(l a) Alpha chain, IgA Bazin et al. (1974) .,S:
;:l
a E5-3 and E5-4 may be the same locus. See references. .,;:l
b Three designations for the rat counterpart of the mouse H-2 locus and human HL·A locus have appeared. ~
, In these two studies, immune response was very closely associated with inheritance of the major histocompatibility locus, and may even be part of a gene complex as is
true for fR-] in mice (McDevitt et al.• 1972).
10. The Laboratory Rat, Rattus norvegicus 249

TABLE 3. Chromosomal Linkage Relationships in the Norway Rat

Chromosomal
designation Gene loci a

LG-I c, Hb-1, Hbb, he, Ig,Jz, p, r, Rw, w

LG-2 an, b, Cu-1, in,s, Sh
LG-3 k, st
LG-4 a,J
LG-5 b Ag-C, Es-1, Es-2, Es-3, Es-4
LG·6 b Ag-B, EAE
LG-7 b GI-1, h

a Gene order of LG-l to 4 is listed as in Robinson (1972) except for the (jz) locus
on LG-l which is now known to be between c (albino) and p (pink-eyed yellow).
b The possibility that LG-5, -6, h-7 may, in fact, be part of LG-3 or -4 has not
been excluded.

which earlier citations are included, or, on occasion, to a particularly rele-

vant review.
The rat has 22 chromosome pairs (Painter, 1928), including the sex
chromosomes X and Y, so the four linkage groups described by Castle
(1947) and the three new groups as described here represent a very small
picture of the rat genome.
The demonstrable linkage of several esterase genes with the Ag-C
blood group locus provided a basis for its designation as LG-5 (Gasser et
at., 19736). Because breeding tests excluding the Ag-C locus from LG-1
and LG-2 have been performed (Palm, 19716, and unpublished), and be-
cause the genes required for establishing a relationship to LG-3 and LG-4
are not known to be available, the use of a new number seems reasonable.
Similarly, two other groups of linked genes may be tentatively designated
as LG-6 and LG-7 (Table 3).
LG-6 includes the Ag-B histocompatibility locus and a gene required
for susceptibility to allergic encephalomyelitis (EAE). These genes appear
to be about 11 recombination units apart, although the exact linkage rela-
tions remain to be determined (Gasser et at., 1973a; Williams and Moore,
1973). LG-7 includes the gene for hooding, h, and production of a plasma
protein, Gt-7 (Moutier et at., 19736).
Table 3 lists the genes associated with both the old and new linkage
groups. Hopefully, this table will receive constant additions and/or revi-
sions in the coming years as the Norway rat, with its many positive at-
tributes for experimental investigation, continues to gain popularity.
Contributing to such progress will be the access of investigators to new
chromosome markers. A considerable number of the mutants listed in
250 O. Mammals

Table 1, indicated by a question mark in the last column, have not been
described for some time and may be lost. This is unfortunate since
renewed interest in the rat has increased the desirability of expanding
knowledge of its genetics. The rapidly increasing number of studies of
antigen-determining loci, studies of immune-response genes, and detec-
tions of gene-controlled and electrophoretic variants, augur well for a new
period of rat genetics.
The need is great for mutant genes which may be used to elaborate
upon the functional regions of specific chromosomes as indicated by their
linkage relationships. Also, with the new techniques for identifying
specific chromosomes (Nilsson, 1973), it should be possible to further
characterize the karyotypic polymorphism exhibited by the rat (Bianchi
and Molina, 1966; Howard, 1971) and its relevance to the expression of
specific mutant loci.
It may be hoped, therefore, that the "missing" mutations of Table 1
will come to the attention of, and kindle a spark of recognition in, investi-
gators in isolated (or nonisolated!) laboratories, so that such mutations
may become generally accessible; new breeding experiments incorporating
old and new mutants should enhance the genetic information, making it
possible to further characterize the biology of the Norway rat, Rattus nor-
vegzGUs.

Acknowledgments

The author is indebted to Ms. Carolyn Rigiero for her able

assistance in developing the list of literature citations. This investigation
was supported, in part, by the United States Publich Health Service re-
search grants CA 10097 and CA 10815 from the National Cancer In-
stitute and RR 05540 from the Division of Research Resources.

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Brdicka, R., 1968 The chromosome I of the laboratory rat, Rattus norvegicus. Acta Univ.
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252 O. Mammals

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10. The Laboratory Rat, Rattus norvegicus 253

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254 O. Mammals

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