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ment of the Kidney, and the Development of the - Journal of Cell ...

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DEVELOPMENT OF AMPHIBIAN KIDNEY 455<br />

controversy that a historical review <strong>of</strong> <strong>the</strong> subject appears to<br />

be justified.<br />

The classic account <strong>of</strong> <strong>the</strong> male urinogenital organs <strong>of</strong> Amphibia<br />

is that <strong>of</strong> Bidder (1846). He first recorded that <strong>the</strong> sperm<br />

traverses <strong>the</strong> kidney <strong>and</strong> noted <strong>the</strong> sperm-duct, lying along <strong>the</strong><br />

medial edge <strong>of</strong> <strong>the</strong> kidney, which to-day bears his name. Many<br />

papers on <strong>the</strong> amphibian urinogenital system appeared during<br />

<strong>the</strong> next thirty years, but none contributed anything to <strong>the</strong><br />

problem <strong>of</strong> <strong>the</strong> genital connexion except a state<strong>ment</strong> by Spengel<br />

(1876) that <strong>the</strong> ordinary excretory tubules have no connexion<br />

with <strong>the</strong> testis. It was left to Nussbaum (1880) to start a controversy<br />

which lasted twenty years.<br />

This author, who appears to have been <strong>the</strong> first to study <strong>the</strong><br />

develop<strong>ment</strong> <strong>of</strong> <strong>the</strong> vasa efferentia, stated categorically that<br />

<strong>the</strong>y were outgrowths from <strong>the</strong> wall <strong>of</strong> <strong>the</strong> malpighian capsule.<br />

These outgrowths grew through <strong>the</strong> mesorchium <strong>and</strong> became<br />

secondarily connected to an independently derived testicular<br />

network. The ducts from <strong>the</strong> kidney were apparent in a twolegged<br />

tadpole, but did not become attached to <strong>the</strong> testicular<br />

network till a few months after metamorphosis. Six years later<br />

<strong>the</strong> same author (Nussbaum, 1886) confirmed Spengel's observation,<br />

which had been made on Eana esculenta while investigating<br />

Eana platyrhinus [=Eana fusca]. In<br />

<strong>the</strong> same year H<strong>of</strong>fman (1886) entered <strong>the</strong> field with <strong>the</strong> story<br />

that every mesonephric unit sent out a connexion to <strong>the</strong> genital<br />

str<strong>and</strong> while this latter was still in <strong>the</strong> undifferentiated condition,<br />

<strong>and</strong> that <strong>the</strong> posterior <strong>of</strong> <strong>the</strong>se kidney-testis connexions<br />

degenerated after metamorphosis. Unfortunately, he also stated<br />

that <strong>the</strong> peritoneal funnels never open into veins (which can be<br />

disproved by <strong>the</strong> examination <strong>of</strong> almost any section—cf. Gray<br />

1930, PL 28, fig. 7), so that his description was generally discounted<br />

on <strong>the</strong> score <strong>of</strong> faulty observation. It appears probable,<br />

never<strong>the</strong>less, that <strong>the</strong> condition he described is <strong>the</strong> primitive<br />

one in Amphibia, for it is very similar to that described by<br />

Semon (1890) for Ichthyophis. In this animal an epi<strong>the</strong>lial<br />

str<strong>and</strong> runs out from <strong>the</strong> wall <strong>of</strong> each mesonephric duct, 'der<br />

sich in zwei Arme gabelt; der eine tritt zur Nebenniere, der<br />

<strong>and</strong>ere zur Keimdriise. Beide sind Derivate der ursprunglichen

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