Saturday, 19 October 2013

Do not marsupials have trophoblast?

Embryonic and trophoblastic areas of the marsupial
blastocyst as envisaged by Hartman 

In the final chapter of her book (previous post), Zofia Kielan-Jaworowska writes on placental mammals, "Their enormous success depends in the first place on the acquisition of the trophoblast, on the basis of which the placenta is formed." In contrast, "Without the trophoblast, marsupials are unable to prolong their gestation period and are born at an extremely early level of anatomical development." This view or a version of it seems to be shared by other zoologists. 

The souce of the misunderstanding can be traced to influential papers by Jason A. Lillegraven (e.g. here). The marsupial blastocyst lacks the inner cell mass of a placental mammal blastocyst. Early on, however, there are two distinct areas that often have been referred to as the embryonic and trophoblastic areas. Lillegraven argues, convincingly in my view, that the "embryonic area" is not equivalent to the inner cell mass and that a considerable portion of the "trophoblastic area" forms part of the developing embryo. In an anatomical sense the trophoblastic area is not the trophoblast as defined by Hubrecht (see previous post).

Nonetheless, Lillegraven also says, "But from a functional point of view (as applied to extra-embryonic parts of the conceptus having direct nutritive significance), the term "trophoblast" may be used to good advantage throughout the Amniota, whether development occurs within a shelled or a shell-free setting."

Thus marsupials do have trophoblast and of course they all have at least a yolk sac placenta and sometimes a chorioallantoic placenta to boot. As shown by Ulrich Zeller and Claudia Freyer (here) there are marsupials where the trophoblast forms  a syncytium and exhibits invasive properties.

As noted in a previous post, there are sound reasons for distinguishing between Eutheria and Placentalia but it is little wonder that marsupial specialists bridle at the term "placental mammals."  

Friday, 4 October 2013

Karl Bogislaus Reichert

Karl Bogislaus Reichert (1811-1883)

Crucial to our understanding of the evolution of mammals is the origin of the bones of the inner ear from what were elements of the jaw articulation in reptiles. This was worked out in 1837 by the German embryologist Karl Bogislaus Reichert. Zofia Kielan-Jaworowska (previous post) notes this is all the more remarkable in view of the primitive tools available at the time. Later Ernst Gaupp built on this work and the supposed origin of the inner ear bones is known as the Reichert-Gaupp theory.

Reichert's membrane (Rm) in the placenta of the capybara
From Kanashiro et al. Reprod Biol Endocrinol 2009; 7: 57 (here)

Students of the placenta will be more familiar with the name of Reichert through the eponymous membrane that lies between the trophoblast and the endoderm of the parietal yolk sac in rodents and insectivores. It was first described by Reichert in the guinea pig (Abh Akad Wissensch Berlin 1862; pp. 97-216).