Wednesday, 28 May 2014

Placentation in a marmot: the woodchuck

Woodchuck or groundhog (Marmota monax)
CC Wikimedia Commons
Not much has been written about placentation in squirrels and other sciuromorph rodents - at least in comparison to myomorphs (e.g. mouse, hamster) and hystricomorphs (e.g. guinea pig, capybara). Marmots belong with chipmunks and ground squirrels in the Tribe Marmotini. The most complete description is for the thirteen-lined ground squirrel (Ictidomys tridecemlineatus) (Mossman & Weisfeldt Am J Anat 1939;64:59-109). It was based largely on specimens collected a century ago by Thomas George Lee (background here).

Interhaemal region in the labyrinth of a woodchuck placenta
Marmota monax Courtesy of Dr. Allen C. Enders
The woodchuck has a labyrinthine, haemochorial placenta. In the above section the large channels with maternal red cells are lined by syncytiotrophoblast. A fetal capillary is seen to the right.

Early Development of the placenta in the Colorado chipmunk
Tamias quadrivittatus Courtesy of Dr. Allen C. Enders
There are several ways to make a haemochorial placenta (discussed here). Squirrels go through a transient endotheliochorial phase. The figure is from a chipmunk and shows a maternal capillary in which part of the endothelium has been replaced by syncytiotrophoblast whilst two endothelial cells remain intact (further figures here). In the first instance this creates the equivalent of the spongy zone found in other rodent placentas.

Later the fetal mesoderm grows into the trophoblast bringing with it the fetal capillaries. To start with the outgrowths are fingerlike (villi) as can be seen in a recent publication on the woodchuck (here). Nearer term, however, the labyrinth occupies most of the depth of the placenta. The spongy zone is then very thin and occupied by syncytiotrophoblast with clumps of nuclei as well as mononucleate giant cells (Dr. Allen C. Enders, personal communication).

Syncytins are endogenous retrovirus envelope genes (previous post). Two occur in murid rodents and another in South American hystricomorphs. Now a search of the genome of the thirteen-lined ground squirrel has turned up several candidates and further work in the woodchuck has shown one of them to be a bona fide syncytin. By in situ hybridization the gene was not expressed in the labyrinth but rather in the part of the spongy zone that had yet to be reached by the fetal vessels.

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