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on the development of the hypobranchial and laryngeal muscles in ...

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158 F. H. Edgeworta<br />

mordium comm<strong>on</strong> to it <strong>and</strong> <strong>the</strong> C<strong>on</strong>strictor oesophagi. The <strong>on</strong>ly muscle which<br />

possibly may be comm<strong>on</strong> to Dipnoi <strong>and</strong> Amphibia is a C<strong>on</strong>strictor laryngis,<br />

which is present <strong>in</strong> Lepidosiren <strong>and</strong> Protopterus1.<br />

In Amniota, as <strong>in</strong> Amphibia, <strong>the</strong> transverse groove is developed beh<strong>in</strong>d<br />

<strong>the</strong> branchial regi<strong>on</strong> <strong>in</strong> <strong>the</strong> floor <strong>of</strong> <strong>the</strong> oesophagus, <strong>and</strong> <strong>the</strong> <strong>laryngeal</strong> groove<br />

is formed progressively forward <strong>in</strong>to <strong>the</strong> br<strong>on</strong>chial regi<strong>on</strong>. The differences<br />

from Amphibia are that <strong>the</strong> <strong>laryngeal</strong> groove extends <strong>in</strong>to a more anterior<br />

br<strong>on</strong>chial segment <strong>and</strong> that <strong>the</strong> separati<strong>on</strong> <strong>of</strong> <strong>the</strong> respiratory epi<strong>the</strong>lium<br />

extends relatively fur<strong>the</strong>r forward so that <strong>the</strong> larynx lies entirely <strong>in</strong> <strong>the</strong><br />

br<strong>on</strong>chial regi<strong>on</strong>. Corresp<strong>on</strong>d<strong>in</strong>gly, <strong>the</strong> primordium <strong>of</strong> <strong>the</strong> <strong>laryngeal</strong> <strong>muscles</strong><br />

separates from that <strong>of</strong> <strong>the</strong> C<strong>on</strong>strictor oesophagi, migrates forward <strong>in</strong>to <strong>the</strong><br />

br<strong>on</strong>chial regi<strong>on</strong> <strong>and</strong> <strong>the</strong>re develops <strong>in</strong>to <strong>the</strong> <strong>laryngeal</strong> <strong>muscles</strong>. This method<br />

is sec<strong>on</strong>dary <strong>and</strong> related to <strong>the</strong> sec<strong>on</strong>dary positi<strong>on</strong> <strong>of</strong> <strong>the</strong> larynx.<br />

In Sauropsida <strong>the</strong> <strong>muscles</strong> c<strong>on</strong>sist <strong>of</strong> a Dilatator <strong>and</strong> a C<strong>on</strong>strictor laryngis<br />

which are homologous with those <strong>of</strong> Amphibia. In a few Reptiles Laryngei<br />

are developed. These, like <strong>the</strong> C<strong>on</strong>strictor, lie posterior to <strong>the</strong> <strong>in</strong>serti<strong>on</strong> <strong>of</strong><br />

<strong>the</strong> Dilatator, are modificati<strong>on</strong>s <strong>of</strong> <strong>the</strong> C<strong>on</strong>strictor <strong>and</strong> not homologous with<br />

<strong>the</strong> Laryngei <strong>of</strong> Urodela <strong>and</strong> Gymnophi<strong>on</strong>a.<br />

In Mammals <strong>the</strong> <strong>laryngeal</strong> <strong>muscles</strong> c<strong>on</strong>sist <strong>of</strong> a Dilatator, Interarytenoid<br />

<strong>and</strong> Laryngeus ventralis. The Dilatator is homologous with that <strong>of</strong> Amphibia<br />

<strong>and</strong> Sauropsida <strong>and</strong>, as <strong>in</strong> <strong>the</strong> latter, arises from <strong>the</strong> Cricoid cartilage. The<br />

Interarytenoid represents <strong>the</strong> dorsal half <strong>of</strong> a C<strong>on</strong>strictor. Both <strong>the</strong> Interarytenoid<br />

<strong>and</strong> Laryngeus ventralis lie <strong>in</strong> fr<strong>on</strong>t <strong>of</strong> <strong>the</strong> <strong>in</strong>serti<strong>on</strong> <strong>of</strong> <strong>the</strong> Dilatator.<br />

Apparently, <strong>the</strong>refore, <strong>the</strong>y are homologous, not with <strong>the</strong> C<strong>on</strong>strictor <strong>of</strong><br />

Sauropsida <strong>and</strong> Amphibia, but with <strong>the</strong> Laryngei <strong>of</strong> Urodela <strong>and</strong><br />

Gymnophi<strong>on</strong>a.<br />

Recurrent <strong>laryngeal</strong> nerve. As <strong>the</strong> <strong>laryngeal</strong> <strong>muscles</strong> are not br<strong>on</strong>chial <strong>in</strong><br />

orig<strong>in</strong>, <strong>the</strong>ir motor nerve-<strong>the</strong> recurrent <strong>laryngeal</strong>-is not a br<strong>on</strong>chial nerve.<br />

It may be regarded as a specialised oesophageal branch <strong>of</strong> <strong>the</strong> vagus. In<br />

Gymnophi<strong>on</strong>a <strong>and</strong> Urodela, <strong>and</strong> markedly <strong>in</strong> <strong>the</strong> latter, it extends to <strong>the</strong><br />

ventral br<strong>on</strong>chial <strong>muscles</strong>, supplement<strong>in</strong>g or supplant<strong>in</strong>g br<strong>on</strong>chial nerves.<br />

The phylogenetic history <strong>of</strong> <strong>the</strong> recurrent <strong>laryngeal</strong> nerve is obscure. No<br />

nerve Was described by v. Wijhe or Beauregard <strong>in</strong> Ceratodus, nor by Hyrtl<br />

<strong>in</strong> Lepidosiren. P<strong>in</strong>kus described, <strong>in</strong> Protopterus, a f<strong>in</strong>e twig extend<strong>in</strong>g from<br />

<strong>the</strong> N. <strong>in</strong>test<strong>in</strong>alis to <strong>the</strong> mucous membrane <strong>of</strong> <strong>the</strong> pharynx <strong>and</strong> larynx, <strong>and</strong><br />

a R. muscularis <strong>and</strong> recurrens-a str<strong>on</strong>g branch from <strong>the</strong> vagus gangli<strong>on</strong><br />

which passes down <strong>on</strong> <strong>the</strong> outside <strong>of</strong> <strong>the</strong> " C<strong>on</strong>strictor pharyngis " <strong>and</strong> divides<br />

<strong>in</strong>to two branches, <strong>on</strong>e <strong>of</strong> which passes forwards <strong>and</strong> s<strong>in</strong>ks <strong>in</strong>to <strong>the</strong> t<strong>on</strong>gue<br />

<strong>muscles</strong>, whilst <strong>the</strong> o<strong>the</strong>r <strong>in</strong>nervates <strong>the</strong> "C<strong>on</strong>strictor pharyngis." This last<br />

branch was also described by Agar. Nei<strong>the</strong>r observer states whe<strong>the</strong>r twigs<br />

from this R. muscularis to <strong>the</strong> C<strong>on</strong>strictor oesophagi can be traced <strong>in</strong>to <strong>the</strong><br />

I It is to be noted that Greil's <strong>in</strong>vestigati<strong>on</strong>s <strong>of</strong> <strong>the</strong> larynx <strong>of</strong> Ceratodus did not extend bey<strong>on</strong>d<br />

<strong>the</strong> 18 mm. stage <strong>and</strong> that Wiedersheim said that <strong>the</strong> state <strong>of</strong> his material left much to be desired.<br />

So it is possible that Ceratodus, too, has a C<strong>on</strong>strictor laryngis.

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