6533b821fe1ef96bd127b76b

RESEARCH PRODUCT

The ontogeny of the filter apparatus of anuran larvae (Amphibia, Anura)

subject

description

The pharynx ofBufo calamita, Rana temporaria andBombina variegata larvae (larval Types IV and III) changes considerably during the latter part of embryonic development. The entodermal regions between the visceral pockets flatten inward to form the anlagen of the filter plates. The ectoderm thrusts forward from the area of the persistent epidermal gills overlying the anlagen of the filter plates. The esophagus pushes dorsolaterally into the pharynx to give rise to the ciliary cushions. Comparison with the development ofXenopus laevis (larval Type I) reveals shared characters: (1) the filter plates are overlapped by the sensory layer of the epiderm and (2) the ciliary grooves are, like the ciliary cushions of larval Types III and IV, anteriorly directed dorsolateral extensions of the esophagus. In all the species studied an ectodermal-esophageal filter apparatus develops. The evolutionary origin of this filter apparatus is discussed. The epidermalization of gills is suggested as a common character with the sister group of Dipnoi, and is therefore a plesiomorphic character in all amphibians. The tendency of filter plate epidermalization is considered to be the end of a process which is also indicated in the epidermalization of the first visceral pouch in lung fish. The ciliary groove is unique in anuran larvae within the Lissamphibia, and is therefore seen as an autapomorphic character within amphibians. On the basis of the different structure of the ciliary cushion inX. laevis and in the other species of this study, two alternative levels of evolutionary ciliary groove origin are discussed. Derivation from the esophagus took place: (1) in a common anuran larval ancestor, or (2) at two independent levels; the first in the Pipidae (-Rhinophrynidae) ancestor and the second in the ancestor of all the other anuran families. Several larval characters and cladistic aspects make the first alternative more probable than the second. Larval Type II anatomy and Larval Type II truncation from the Larval Type IV of Ranoidea do not contradict these considerations. There is disproportionately early commencement of ingestion activity inR. temporaria (G Stage 23),B. calamita (G Stage 23), andB. bufo (G Stage 24) compared toXenopus. Feeding in the former three species precedes the differentiation of the filter plates, their mucus production, and the exhaustion of the yolk supply in the gut tissue. By contrast, the goblet cells and the ciliary cells of the ciliary cushions are already differentiated when feeding starts. This suggests that ingestion in these early stages requires mucus production by the ciliary cushions and transport by their ciliary cells. Presumably in fully formed larvae, the ciliary cushions are the mucus donors, whereas the filter plates are the mucus depositors. By contrast,X. laevis does not begin active food intake by suspension feeding until after the yolk supply has been used up from the entoderm of the buccal cavity to deep in the esophagus.