, the bursa of Fabricius
: Bursa cloacalis
or Bursa fabricii
) is the site of hematopoiesis
, a specialized organ
that, as first demonstrated by Bruce Glick
and later by Max Cooper
and Robert Good
, is necessary for B cell
development in birds. Mammals
generally do not have an equivalent organ; the bone marrow
is often both the site of hematopoiesis and B cell development.
It is named after Hieronymus Fabricius.
The ‘B’ in ‘B cell’ refers to bursa-derived. This is simply because during the 1960s B cells were first
defined (and distinguished from thymus-derived T cells) in birds, which have a bursa. A decade later, after examining almost every other organ including the appendix, researchers finally discovered that mammalian B cells develop in the bone marrow and spleen. The fact that ‘bone marrow’, like bursa, starts with a ‘B’ is a coincidence.
The bursa is an epithelial and lymphoid organ that is found only in birds. The bursa develops as a dorsal diverticulum of the proctadael region of the cloaca. The luminal surface of the bursa is plicated with as many as 15 primary and 7 secondary plicae or folds. These plicae have hundreds of bursal follicles containing follicle-associated epithelial cells, lymphocytes, macrophages, and plasma cells. Lymphoid stem cells migrate from the fetal liver to the bursa during ontogeny. In the bursa, these stem cells acquire the characteristics of mature, immunocompetent B cells.
In 1956, Bruce Glick showed that removal of the bursa in newly hatched chicks severely impaired the ability of the adult birds to produce antibodies. In contrast, removal of the bursa in adult chickens has not much effect on the immune system. This was a serendipitous discovery that came about when a fellow graduate, Timothy S. Chang, who was teaching a course on antibody production obtained chickens from Glick that had been bursectomised (removal of the bursa). When these chickens failed to produce antibody in response to an immunization with Staphylococcus
bacteria, the two students realized that the bursa is necessary for antibody production. While some have suggested that this observation was worthy of a Nobel prize
, none was awarded, and in fact their initial attempts to publish their findings were thwarted by an editor who commented that "further elucidation of the mechanism ... should be attempted before publication.”
The role of the thymus in the immune response was also identified shortly after the discovery of bursa’s role in antibody responses. In thymectomized animals, the ability to reject allografts, and to mount delayed hypersensitivity responses, was drastically reduced. By the mid-1960s, immunologists were convinced that there were indeed two separate arms of the immune system: one dealing exclusively with the production of circulating antibodies (humoral immunity), and another that is involved in the delayed hypersensitivity-type reactions and graft rejections (cell-mediated immunity).