Rhesus Macaque[1] | |
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Conservation status | |
Least Concern (IUCN 3.1)[2] |
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Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Family: | Cercopithecidae |
Genus: | Macaca |
Species: | M. mulatta |
Binomial name | |
Macaca mulatta (Zimmermann, 1780) |
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Rhesus Macaque range |
The Rhesus Macaque (Macaca mulatta), often called the Rhesus Monkey, is one of the best known species of Old World monkeys.
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Adult males measure approximately 53 centimeters on average and weigh an average of 7.7 kilograms. Females are smaller, averaging 47 centimeters in length and 5.3 kilograms in weight. This macaque is brown or grey in color and have pink faces which are typically bereft of fur. Its tail is of medium length and averages between 20.7 and 22.9 centimeters. It typically has a lifespan of about 25 years.
The species is native to northern India, Bangladesh, Pakistan, Nepal, Burma, Thailand, Afghanistan, Vietnam, southern China, and some neighboring areas. The rhesus monkey has the widest geographic ranges of any nonhuman primate, occupying a great diversity of altitudes throughout Central, South, and Southeast Asia. Inhabiting arid, open areas, the rhesus macaques may be found in grasslands, woodlands and in mountainous regions up to 2,500 metres in elevation. They are regular swimmers. Babies as young as a few days old can swim, and adults are known to swim over a half mile between islands, but are often found drowned in small groups where their drinking waters lie. Rhesus macaques are noted for their tendency to move from rural to urban areas, coming to rely on handouts or refuse from humans.[3] They have become a pest in some areas, perceived as a possible risk to public health and safety.
A diurnal animal, the Rhesus macaque is both arboreal and terrestrial, mostly herbivorous feeding on leaves and pine needles, roots, and the occasional insect or small animal. The monkey has specialized pouch-like cheeks, allowing it to temporarily hoard its food.
The name rhesus is reminiscent of the Greek mythological king Rhêsos. However, the French naturalist Jean-Baptiste Audebert who applied the name to the species, stated that it had no meaning.[4]
According to Zimmermann’s first description of 1780, the rhesus macaque is distributed in eastern Afghanistan, Bangladesh, Bhutan, as far east as the Brahmaputra Valley in peninsular India, Nepal and northern Pakistan. Today, this is known as the Indian rhesus macaque Macaca mulatta mulatta, which includes the morphologically similar Macacus rhesus villosus described by True in 1894 from Kashmir and Macaca mulatta mcmahoni described by Pocock in 1932 from Kootai, Pakistan. Several Chinese subspecies of rhesus macaques have been described between 1867 and 1917. The molecular differences identified among populations however are alone not consistent enough to conclusively define any subspecies, but only grade C taxa requiring further investigation.[5]
The Chinese subspecies can be divided in:
Around the spring of 1938, a colony of rhesus macaques were released in the Silver River State Park in Florida by a tour boat operator known locally as "Colonel Tooey" to enhance his "Jungle Cruise". A traditional story that the monkeys were released for scenery enhancement in the Tarzan movies that were filmed at that location is false, as the only Tarzan movie filmed in the area, 1939's Tarzan Finds a Son! contains no rhesus macaques.[8] In addition, various colonies of rhesus and other monkey species are speculated to be the result of zoos and wildlife parks destroyed in hurricanes, most notably Hurricane Andrew.[9]
There is also a notable colony of rhesus macaques on Morgan Island, one of the Sea Islands in the South Carolina Lowcountry. They were imported in the 1970s for use in local labs and are by all accounts thriving.[10][11]
Like other macaques, the Rhesus troop comprises a mixture of males and females. The troop may contain up to 180 individuals, but 20 is the average. Females may outnumber the males by a ratio of 4:1. The social hierarchy is also matriarchal, rank dependent on lineage to the lead female. Care of young and territory surveillance duties are shared amongst the troop. While females are more or less placid, males are typically rowdy between themselves. The Rhesus Macaque is characterised as a vociferous monkey. Monkeys that discover food will normally advertise the fact by specific calls, though it has been claimed that young or subordinate monkeys will sometimes seek to avoid doing so if their discovery has gone unobserved. Females cycle similar to humans with menstrual cycles of around 28 days.[12]
Mating is not confined to a specific season. Gestation may last 164 days. Females are mature by three years of age, and males at four. The typical lifespan of a rhesus monkey in captivity is approximately 15–20 years for males and 20–25 years for females.[12] These monkeys rarely live beyond 15 years of age in the wild.
They have a reputation as aggressive urban pests. Their penchant for thievery and aggression can make them somewhat dangerous, particularly to small children.
The Rhesus Macaque is well known to science owing to its relatively easy upkeep in captivity, and has been used extensively in medical and biological research. It has given its name to the Rhesus factor, one of the elements of a person's blood group, by the discoverers of the factor, Karl Landsteiner and Alexander Wiener. The Rhesus Macaque was also used in the well-known experiments on maternal deprivation carried out in the 1950s by controversial comparative psychologist Harry Harlow.
The U.S. Army, the U.S. Air Force, and NASA launched Rhesus Macaques into outer space during the 1950s and 60s, and the Soviet/Russian space program launched them into space as recently as 1997 on the Bion missions. One of these primates ("Able") who was launched on a suborbital spaceflight in 1959 was one of the two first living beings (along with "Miss Baker" on the same mission) to travel in space and return alive.
In January 2000, the Rhesus Macaque became the first cloned primate with the birth of Tetra. January 2001 saw the birth of ANDi, the first transgenic primate; ANDi carries foreign genes originally from a jellyfish.
Work on the genome of the Rhesus Macaque was completed in 2007,[13] making Rhesus Macaque the second non-human primate to have its genome sequenced. The study [14] claims that humans and macaques share about 93% of their DNA sequence and shared a common ancestor roughly 25 million years ago.
Though most studies of the Rhesus Macaque are from various locations in northern India, some knowledge of the natural behavior of the species comes from studies carried out on a colony established by the Caribbean Primate Research Center of the University of Puerto Rico on the island of Cayo Santiago, off Puerto Rico. There are no predators on the island, and humans are not permitted to land except as part of the research programmes. The colony is provisioned to some extent, but about 50% of its food comes from natural foraging. In other more controlled settings, these macaques often enjoy Fig Newtons, and are particularly keen on "pouching" large quantities of marshmallow.
Comparison of rhesus macaques, chimpanzees and humans revealed the structure of ancestral primate genomes, positive selection pressure and lineage-specific expansions and contractions of gene families.
"The goal is to reconstruct the history of every gene in the human genome," said Evan Eichler, University of Washington, Seattle. DNA from different branches of the primate tree will allow us "to trace back the evolutionary changes that occurred at various time points, leading from the common ancestors of the primate clade to Homo sapiens," said Bruce Lahn, University of Chicago.[15]
After the human and chimpanzee genomes were sequenced and compared, it was usually impossible to tell whether differences were the result of the human or chimpanzee gene changing from the common ancestor. After the Rhesus macaque genome was sequenced, 3 genes could be compared. If 2 genes were the same, they are presumed to be the original gene.[16]
The chimpanzee and human genome, which diverged 6 million years ago, had 98% identity and many conserved regulatory regions. Comparing the macaque and human genome, which diverged 25 million years ago and had 93% identity, further identified evolutionary pressure and gene function.
Like the chimpanzee, changes were on the level of gene rearrangements rather than single mutations. There were frequent insertions, deletions, changes in the order and number of genes, and segmental duplications near gaps, centromeres and telomeres. So macaque, chimpanzee and human chromosomes are mosaics of each other.
Surprisingly, some normal gene sequences in healthy macaques and chimpanzees cause profound disease in humans. For example, the normal sequence of phenylalanine hydroxylase in macaques and chimpanzees is the mutated sequence responsible for phenylketonuria in humans. So humans must have been under evolutionary pressure to adopt a different mechanism.
Some gene families are conserved or under evolutionary pressure and expansion in all 3 primate species, while some are under expansion uniquely in human, chimpanzee or macaque.
For example, cholesterol pathways are conserved in all 3 species (and other primate species). In all 3 species, immune response genes are under positive selection, and genes of T cell-mediated immunity, signal transduction, cell adhesion, and membrane proteins generally. Genes for keratin, which produce hair shafts, were rapidly evolving in all 3 species, possibly because of climate change or mate selection. The X chromosome has 3 times more rearrangements than other chromosomes. The macaque gained 1,358 genes by duplication.
Triangulation of human, chimpanzee and macaque sequences showed expansion of gene families in each species.
The PKFP gene, important in sugar (fructose) metabolism, is expanded in macaques, possibly because of their high-fruit diet. So are genes for the olfactory receptor, cytochrome P450 (which degrades toxins), and CCL3L1-CCL4 (associated in humans with HIV susceptibility).
Immune genes are expanded in macaques, relative to all 4 great ape species. The macaque genome has 33 major histocompatibility genes, 3 times that of human. This has clinical significance because the macaque is used as an experimental model of the human immune system.
In humans, the PRAME (preferentially expressed antigen of melanoma) gene family is expanded. It is actively expressed in cancers but normally testis-specific, possibly involved in spermatogenesis. The PRAME family has 26 members on human chromosome 1. In the macaque, it has 8, and has been very simple and stable for millions of years. The PRAME family arose in translocations in the common mouse-primate ancestor 85 million years ago, and is expanded on mouse chromosome 4.
Agilent and Affymetrix have macaque DNA microarrays with 20,000 gene sequences, and they are used in macaque research. For example, Michael Katze of University of Washington, Seattle, infected macaques with 1918 and modern influenza. The DNA microarray showed the macaque genomic response to human influenza on a cellular level in each tissue. Both viruses stimulated innate immune system inflammation, but the 1918 flu stimulated stronger and more persistent inflammation, causing extensive tissue damage, and it did not stimulate the interferon-1 pathway. The DNA response showed a transition from innate to adaptive immune response over 7 days.
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