Biomedical Engineering Reference
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Indochina where the two antigens are approximately
equally prevalent in both species ( Malaivijitnond et al.,
2008 )]; the O antigen is present, but rare, in both species.
Farkas et al. (2010) reported that 97% of 500 Indian rhesus
macaques from the Tulane National Primate Research
Center were blood type B by detecting agglutinins in saliva.
Unfortunately, regional variation in the distribution of ABO
phenotyes among populations of neither species has been
well-characterized using serological tests. The similarity of
allele frequencies of the two species in Indochina, but not
elsewhere, is consistent with the hypothesis that the two
species have naturally interbred extensively in Indochina.
Some of the studies cited above reported the presence of
blood group O. However, the failure to absorb the NHP sera
with human type O RBCs may have caused nonspecific
agglutination of species-specific agglutinins to human
A and B test cells as demonstrated by Chen et al. (2009) ,
who also found B to predominate in rhesus and cynomolgus
macaques and A to be more common in cynomolgus
macaques than previously reported when sera were not
absorbed with human type O RBCs.
A multiplex PCR test was developed that amplifies the
region containing the codons responsible for the A and B
antigens and found rhesus macaques to predominantly
carry the B with a moderate frequency of the A allele and
an absence of the O allele. More significantly for trans-
plantation research using nonhuman primates, the random
probability of major ABO incompatibility exceeded 40%
( Premasthusan et al., 2011 ). Thus, it is paramount that
nonhuman primates used in transplantation studies, most of
whom express the ABH substances, be reliably phenotyped
for their ABO phenotype.
polymorphisms ( Kanthaswamy and Smith, 1997; Morin
et al., 1997; Smith et al., 2000, 2006; Smith, 2005 ),
Mhc alleles ( Viray et al., 2001; Doxiadis et al., 2003 ;
Sauermann, personnel communication), and mtDNA
( Melnick et al., 1993; Kanthaswamy and Smith, 2004;
Smith and McDonough, 2005; Smith et al., 2006, 2007;
Satkoski et al., 2008a; Stevison and Kohn, 2008 ). These
studies add support to the proposal that at least two, one
eastern and one western, subspecies of rhesus macaque be
recognized. Studies of Indochinese rhesus macaques are
needed to determine whether or not
they also warrant
a separate subspecies distinction.
mtDNA
Differences in mtDNA are particularly useful in the study
of intra-species variation because they evolve rapidly
( Brown et al., 1979 ) and are solely maternally inherited
( Giles et al., 1980 ), which precludes recombination. The
former trait allows differentiation among relatively closely
related individuals (i.e. intraspecific differences) while the
latter permits mtDNA haplotypes to be arranged in ances-
ter/descendant relationships. Its maternal inheritance also
fosters regional diversity in mtDNA, because most Old
World monkeys exhibit female philopatry (i.e. females
remain and breed in their natal groups while males disperse
after reaching sexual maturity ( Pusey and Packer, 1987 )),
leading to marked geographical structure to mtDNA vari-
ation. However, mtDNA, like Y-chromosome haplotypes, is
effectively a single locus and so is subject to more
stochastic processes of evolution due to its lower effective
population size than nuclear DNA and sometimes provides
evidence contradictory to that provided by nuclear genes
( Evans et al., 2003 ).
Since every gene can experience a unique evolutionary
history and exhibit a correspondingly unique gene tree,
phylogenetic and phylogeographical studies of primate
species should employ multiple loci that have experienced
a neutral (nonadaptive) evolutionary history. In general, the
number of loci employed in such studies is much more
important than the number of samples of any given species
included in the study as long as the sample size allows
discovery of all but the rarest variants that contribute little
to estimates of gene diversity.
Microsatellite Polymorphisms
Later, microsatellite (STR) polymorphisms, which are far
more informative than protein coding or blood group
polymorphisms, with gene diversity values typically
exceeding 0.7, were identified in rhesus and longtail
macaques by using the polymerase chain reaction (PCR)
cross-amplification with human primers ( Ely et al., 1996,
1999; Kayser et al., 1996; Rogers et al., 2005, 2006;
Kanthaswamy et al., 2006; Kikuchi et al., 2007 ). Variation
in frequencies of Mhc alleles ( Bontrop, 1994; Bontrop
et al., 1996; Sauermann 1996, 1998 ), mitochondrial DNA
(mtDNA) restriction site haplotypes (e.g. Melnick et al.,
1993; Zhang and Shi, 1993 ) and sequences ( Hayasaka
et al., 1996; Li and Zhang, 2005 ), functional genes
( Deinard and Smith, 2001 ), and Y-chromosome haplotypes
( Tosi et al., 2000 ) have also been characterized. Since then,
additional genetic differences between populations of rhe-
sus macaques from China and India have been reported that
are based on electrophoretically defined protein poly-
morphisms ( Fooden and Lanyon, 1989; Smith, 1994 ), STR
Macaques
The above studies report levels of genetic heterogeneity
that are, as in the study of Nozawa et al. (1977) , variable but
approximately equal, on average, between rhesus and
longtail macaques, higher in rhesus macaques of Chinese
than in those of Indian origin (suggesting major differences
in past demographical experiences of the two regional
populations), and consistently lower in M. fuscata,
M. radiatta, and certain isolated island populations of
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