Absence of mitral cells in monolayer in monotremes. Variations in vertebrate olfactory bulbs.

An invariant feature of the olfactory bulb in placental and marsupial mammals is the arrangement of the perikarya of mitral cells in a monolayer. Contrasting with this is the arrangement found in the olfactory bulbs of the monotremes, platypus and echidna, where the large perikarya are not only absent from the position of a monolayer (usually forming the external boundary of the internal plexiform layer) but occupy a region which would characterize them as tufted cells. In other classes of amniote vertebrates, reptiles and birds, the placement of large perikarya in the olfactory bulb ranges from a compact layer to a broad band. Such an overview among several vertebrate classes suggests that a monolayer of mitral cells may be a specialized subset of the tufted-mitral cell population. The accessory olfactory formation among mammals also exhibits variation in the compactness of the large perikarya: a broadband in most but a compact layer in a few others such as the chinchilla and the capybara. Such specialized alignment of perikarya (and, consequently, of their dendritic and axonal elements) may enable more refined signal processing than does random alignment of these elements. Such speculations can be tested using appropriate phylogenetic sampling, and monotremes provide particularly advantageous test cases.