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156 F. H. Edgeworth<br />

The <strong>development</strong> <strong>of</strong> <strong>the</strong> pars trachealis cartilag<strong>in</strong>is lateralis is related to<br />

<strong>the</strong> formati<strong>on</strong> <strong>of</strong> a trachea which develops at <strong>the</strong> end <strong>of</strong> <strong>the</strong> larval stage <strong>in</strong><br />

Rana, dur<strong>in</strong>g larval life <strong>in</strong> Menopoma, Amblystoma, Trit<strong>on</strong>, Siph<strong>on</strong>ops<br />

<strong>and</strong> Ichthyophis, <strong>and</strong> <strong>in</strong> <strong>the</strong> 20 mm. stage <strong>of</strong> Necturus. The pars trachealis<br />

is formed as a n<strong>on</strong>-ch<strong>on</strong>drified backward prol<strong>on</strong>gati<strong>on</strong> <strong>of</strong> <strong>the</strong> pars laryngea<br />

s. arytenoid. It may persist <strong>in</strong> this c<strong>on</strong>diti<strong>on</strong>, e.g. Necturus, or may subsequently<br />

ch<strong>on</strong>drify <strong>in</strong> tracts <strong>of</strong> more or less complete r<strong>in</strong>gs. In Trit<strong>on</strong> <strong>and</strong><br />

Salam<strong>and</strong>ra <strong>the</strong> cartilage is first developed at <strong>the</strong> bifurcati<strong>on</strong> <strong>of</strong> <strong>the</strong> trachea.<br />

The above discussi<strong>on</strong> shows that fur<strong>the</strong>r embryological <strong>in</strong>vestigati<strong>on</strong>s are<br />

needed-<strong>in</strong> particular it would be well to know <strong>the</strong> c<strong>on</strong>diti<strong>on</strong> <strong>of</strong> <strong>the</strong> br<strong>on</strong>chial<br />

skelet<strong>on</strong> <strong>and</strong> ventral br<strong>on</strong>chial <strong>muscles</strong> <strong>in</strong> <strong>the</strong> larvae <strong>of</strong> Amphiumal <strong>and</strong><br />

Cryptobranchus jap., <strong>the</strong> <strong>development</strong> <strong>of</strong> <strong>the</strong> sph<strong>in</strong>cter <strong>laryngeal</strong> fibres <strong>in</strong><br />

Siren, <strong>and</strong> <strong>the</strong> <strong>development</strong> <strong>of</strong> <strong>the</strong> ventral br<strong>on</strong>chial <strong>muscles</strong>, larynx <strong>and</strong><br />

<strong>laryngeal</strong> <strong>muscles</strong> <strong>in</strong> Gymnophi<strong>on</strong>a.<br />

On <strong>the</strong> phylogenetic history <strong>of</strong> <strong>the</strong> larynx, trachea, <strong>and</strong> <strong>laryngeal</strong> <strong>muscles</strong><br />

Schmidt (1913), who <strong>in</strong>vestigated <strong>the</strong> <strong>development</strong> <strong>of</strong> <strong>the</strong> larynx <strong>in</strong><br />

certa<strong>in</strong> Reptiles, was <strong>of</strong> op<strong>in</strong>i<strong>on</strong> that <strong>the</strong> lungs <strong>of</strong> Polypterus, Amphibia, <strong>and</strong><br />

Amniota, like <strong>the</strong> swimm<strong>in</strong>g bladder <strong>of</strong> Lepidosteus <strong>and</strong> Amia, develop <strong>in</strong> <strong>the</strong><br />

same native soil ("Mutterboden"), <strong>and</strong> that, <strong>on</strong> <strong>the</strong> o<strong>the</strong>r h<strong>and</strong>, <strong>the</strong> trachea<br />

with <strong>the</strong> larynx <strong>of</strong> Amniota, <strong>and</strong> possibly also that <strong>of</strong> Amphibia, are phylogenetically<br />

a later formati<strong>on</strong>, <strong>the</strong> <strong>development</strong> <strong>of</strong> which beg<strong>in</strong>s with pulm<strong>on</strong>ary<br />

respirati<strong>on</strong>.<br />

The larynx <strong>of</strong> Dipnoi <strong>and</strong> Polypterus lies <strong>in</strong> <strong>the</strong> ventral wall <strong>of</strong> <strong>the</strong> gut<br />

beh<strong>in</strong>d <strong>the</strong> br<strong>on</strong>chial regi<strong>on</strong>. Its musculature was described by Wiedersheim<br />

(1904) <strong>and</strong> Gbppert (1904). They came to <strong>the</strong> c<strong>on</strong>clusi<strong>on</strong>, though without any<br />

embryological evidence, that it is developed from pharyngeal musculature<br />

<strong>and</strong> that it represents musculature <strong>of</strong> atrophied br<strong>on</strong>chial arches.<br />

Neumeyer (1904) <strong>and</strong> Kellicott (1905), <strong>in</strong>dependently, showed that <strong>the</strong><br />

larynx <strong>of</strong> Ceratodus is developed <strong>in</strong> <strong>the</strong> ventral wall <strong>of</strong> <strong>the</strong> gut at some distance<br />

beh<strong>in</strong>d <strong>the</strong> br<strong>on</strong>chial regi<strong>on</strong>. Greil (1913) c<strong>on</strong>firmed <strong>the</strong>ir observati<strong>on</strong>s, <strong>and</strong><br />

also showed that <strong>the</strong> free mesoderm cells round <strong>the</strong> gut <strong>and</strong> larynx become<br />

sp<strong>in</strong>dle-shaped <strong>and</strong> develop <strong>in</strong>to smooth muscle cells, which form a mantle<br />

round <strong>the</strong> gut <strong>and</strong> larynx.<br />

Kerr (1910) stated that <strong>in</strong> Lepidosiren <strong>and</strong> Protopterus <strong>the</strong> lung-rudiment<br />

is developed, <strong>in</strong> stage 32, as a mid-ventral bulg<strong>in</strong>g from <strong>the</strong> pharynx at <strong>the</strong><br />

level <strong>of</strong> cleft vi. His figures show that its c<strong>on</strong>necti<strong>on</strong> with <strong>the</strong> gut migrates<br />

backwards, so that <strong>in</strong> stage 35 it is dist<strong>in</strong>ctly posterior to <strong>the</strong> level <strong>of</strong> <strong>the</strong><br />

6th gill-clefts, i.e. is <strong>in</strong> <strong>the</strong> oesophageal regi<strong>on</strong>. In Lepidosiren, Agar (1907)<br />

stated that <strong>the</strong> ventral <strong>and</strong> lateral parts <strong>of</strong> <strong>the</strong> "C<strong>on</strong>strictor pharyngis" <strong>of</strong><br />

Wiedersheim are formed from mesenchymatous cells, budded <strong>of</strong>f from <strong>the</strong><br />

<strong>in</strong>ner walls <strong>of</strong> <strong>the</strong> pericardio-perit<strong>on</strong>eal ducts, <strong>and</strong> are thus <strong>of</strong> splanchnic<br />

orig<strong>in</strong>. The dorsal part <strong>of</strong> <strong>the</strong> muscle is <strong>of</strong> somatic orig<strong>in</strong>, be<strong>in</strong>g derived<br />

I In larvae younger than <strong>the</strong> 45 mm. stage <strong>in</strong>vestigated by Hay.

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