THE BLOOD QROUPS O F MARINE ANIMALS
97
including one (not known to vary individually) that has a specificity
close to that of human B antigen. These findings are reported in a
scries of papers by Fujino and his associates, as well as in some
additional papers by Cushing et aE. (references below).
Attention has ccntered upon three phenotypes, J u l , Ju2 and
Jul.2, first discovered in the finback whale Balaenoptera physalus (L.)
(Fujino, 1966), and subsequently in the humpback whale Megapteru
ltodosa (Bonnat.) (Cushing, et al., 19G9), pygmy-blue whale Balaenoptera
musculus L. (Fujino, 1962), sperm whale Physeter ca.lodon (L.) (Cushing
et al., 1963a) and sei whale (Balaenqtera borealis (Lcsson) (Cushing and
Calaprice, N. L., unpubl. data). These phenotypcs appear to be
controlled in finbacks by a major pair of allelic genes, inasmuch as
extensive blood typing has not disclosed any individuals lacking both
of these antigens. A series of a t least three subtypes of the Ju2 antigen
occur, which increases the potential total number of alleles involved.
Isomtibodies reactive with Ju2 are found in J u l , and in Jul.2, Ju2
phenotypes depending upon their Ju2 subtype. A number of species
of domestic animals have natural agglutinins in their serums that
react with the Ju2 antigen, and hetcroimmune serums are easily
induced in rabbitu and chickcns. A similar situation exists in the
humpback, and also appears to hold for the other whales where J u
antigens are known. However, none of thesc species have been as
extensively studied ria the finback. To date it has not been possible to
show Hardy-Weinberg equilibrium relations among the major phenotypes of any sample of North Pacific finbacks on a simple two-allele
hypothesis, the heterozygous class usually being less then expected.
Fujino (1960) has advanced the hypothesis that at least three alleles
are involved, one determining J u l , a second codominant with that
for J u l and determining Ju2 and a third for a Ju2 that is recessive
to J u l . As Fujino later states (1963) this formal hypothesis while
useful may come to be altered particularly with reference to the
varying ef€ects of dosage among the Ju2 subtypes (which for example
could cause a situation like that reported for the A, B type or the
MN type in humans. Cf. Weiner and Wexler, 1958).
Three comparisons have been made of the frequencies of J u
Phenotypes of whales from different areas. The first and most cxtensive
Was of finhack populations in the North Pacific (Fujino, 1960). Here
variations in blood group frequencies werc found to agrce with extensive
marking, morphometric and other data in supporting the conacpt that
North Pacific finbacks are divided into a t least three subpopulations.
One of these forms an isolated unit in the East China Sea, the other
two comprise Eastern ” and ‘ I Western ” Pacific groups which mix
97
including one (not known to vary individually) that has a specificity
close to that of human B antigen. These findings are reported in a
scries of papers by Fujino and his associates, as well as in some
additional papers by Cushing et aE. (references below).
Attention has ccntered upon three phenotypes, J u l , Ju2 and
Jul.2, first discovered in the finback whale Balaenoptera physalus (L.)
(Fujino, 1966), and subsequently in the humpback whale Megapteru
ltodosa (Bonnat.) (Cushing, et al., 19G9), pygmy-blue whale Balaenoptera
musculus L. (Fujino, 1962), sperm whale Physeter ca.lodon (L.) (Cushing
et al., 1963a) and sei whale (Balaenqtera borealis (Lcsson) (Cushing and
Calaprice, N. L., unpubl. data). These phenotypcs appear to be
controlled in finbacks by a major pair of allelic genes, inasmuch as
extensive blood typing has not disclosed any individuals lacking both
of these antigens. A series of a t least three subtypes of the Ju2 antigen
occur, which increases the potential total number of alleles involved.
Isomtibodies reactive with Ju2 are found in J u l , and in Jul.2, Ju2
phenotypes depending upon their Ju2 subtype. A number of species
of domestic animals have natural agglutinins in their serums that
react with the Ju2 antigen, and hetcroimmune serums are easily
induced in rabbitu and chickcns. A similar situation exists in the
humpback, and also appears to hold for the other whales where J u
antigens are known. However, none of thesc species have been as
extensively studied ria the finback. To date it has not been possible to
show Hardy-Weinberg equilibrium relations among the major phenotypes of any sample of North Pacific finbacks on a simple two-allele
hypothesis, the heterozygous class usually being less then expected.
Fujino (1960) has advanced the hypothesis that at least three alleles
are involved, one determining J u l , a second codominant with that
for J u l and determining Ju2 and a third for a Ju2 that is recessive
to J u l . As Fujino later states (1963) this formal hypothesis while
useful may come to be altered particularly with reference to the
varying ef€ects of dosage among the Ju2 subtypes (which for example
could cause a situation like that reported for the A, B type or the
MN type in humans. Cf. Weiner and Wexler, 1958).
Three comparisons have been made of the frequencies of J u
Phenotypes of whales from different areas. The first and most cxtensive
Was of finhack populations in the North Pacific (Fujino, 1960). Here
variations in blood group frequencies werc found to agrce with extensive
marking, morphometric and other data in supporting the conacpt that
North Pacific finbacks are divided into a t least three subpopulations.
One of these forms an isolated unit in the East China Sea, the other
two comprise Eastern ” and ‘ I Western ” Pacific groups which mix