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

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142 P. H. Edgeworth<br />

Ellipsoglossa. My observati<strong>on</strong>s show that it is also present <strong>in</strong> <strong>the</strong> larval stages<br />

<strong>of</strong> Ellipsoglossa <strong>and</strong> Menopoma.<br />

It is a muscle that takes orig<strong>in</strong> from <strong>the</strong> fourth br<strong>on</strong>chial bar, passes<br />

forward <strong>and</strong> separates <strong>in</strong>to three slips (a, b, c) which are attached respectively<br />

to <strong>the</strong> first, sec<strong>on</strong>d <strong>and</strong> third br<strong>on</strong>chial bars. In Siren <strong>on</strong>ly (a) is present.<br />

In Necturus <strong>and</strong> Proteus <strong>the</strong>re is a similar Subarcualis rectus iii which takes<br />

orig<strong>in</strong> from <strong>the</strong> third br<strong>on</strong>chial bar, <strong>and</strong> is attached, (a) to <strong>the</strong> first, <strong>and</strong> (b)<br />

to <strong>the</strong> sec<strong>on</strong>d, br<strong>on</strong>chial bar.<br />

The forward extensi<strong>on</strong> <strong>of</strong> Subarcualis rectus iv <strong>in</strong> fr<strong>on</strong>t <strong>of</strong> <strong>the</strong> third<br />

br<strong>on</strong>chial bar-to <strong>the</strong> sec<strong>on</strong>d <strong>and</strong> first bars-is related to <strong>the</strong> sec<strong>on</strong>dary<br />

functi<strong>on</strong> <strong>and</strong> positi<strong>on</strong> <strong>of</strong> Subarcuales ii <strong>and</strong> iii. (It is noticeable that this<br />

sec<strong>on</strong>dary form <strong>of</strong> Subarcualis rectus iv also exists <strong>in</strong> Amphiuma (adult), <strong>and</strong><br />

that <strong>in</strong> Megalobatrachus max. (adult, with <strong>on</strong>ly two br<strong>on</strong>chial bars) though no<br />

Subarcualis rectus iv is present yet Subarcuales obliqui ii <strong>and</strong> iii <strong>and</strong> not<br />

Subarcuales recti ii <strong>and</strong> iii are present videe Druner). These facts form<br />

additi<strong>on</strong>al evidence <strong>in</strong> favour <strong>of</strong> <strong>the</strong> <strong>the</strong>ory, suggested above, that a Urobranchiale<br />

exists <strong>in</strong> <strong>the</strong> larval forms <strong>of</strong> <strong>the</strong>se Urodela also.)<br />

There are thus five closely related phenomena <strong>in</strong> Urodela. (1) Backward<br />

growth <strong>of</strong> <strong>the</strong> h<strong>in</strong>d end <strong>of</strong> <strong>the</strong> Genio-hyoid ventral to <strong>the</strong> forepart <strong>of</strong> <strong>the</strong><br />

Rectus cervicis. (2) Formati<strong>on</strong> <strong>of</strong> a Urobranchiale. (3) Formati<strong>on</strong> <strong>of</strong> Subarcuales<br />

obliqui ii <strong>and</strong> iii (Subarcualis obliquus ii <strong>in</strong> Necturus <strong>and</strong> Proteus).<br />

(4) Forward extensi<strong>on</strong> <strong>of</strong> Subarcualis rectus iv (Subarcualis rectus iii <strong>in</strong> Necturus<br />

<strong>and</strong> Proteus) to <strong>the</strong> first branchial bar. (5) Separati<strong>on</strong> <strong>of</strong> a superficial,<br />

ventral, porti<strong>on</strong> <strong>of</strong> <strong>the</strong> Rectus cervicis, attached to <strong>the</strong> Urobranchiale.<br />

No <strong>on</strong>e <strong>of</strong> <strong>the</strong>se sec<strong>on</strong>dary phenomena occur <strong>in</strong> Anuran <strong>and</strong> Gymnophi<strong>on</strong>an<br />

larvae, where <strong>the</strong> c<strong>on</strong>diti<strong>on</strong>s are more primitive. (There is a small overlap<br />

<strong>of</strong> <strong>the</strong> Genio-hyoid <strong>and</strong> Rectus cervicis <strong>in</strong> Rana temp. larvae, but it is due to<br />

a backward extensi<strong>on</strong> <strong>of</strong> <strong>the</strong> former al<strong>on</strong>g <strong>the</strong> <strong>hypobranchial</strong> plate, <strong>and</strong> so <strong>of</strong><br />

a different character from that <strong>of</strong> Urodela. The anterior end <strong>of</strong> <strong>the</strong> Rectus<br />

becomes attached to <strong>the</strong> Crista hyoidea (<strong>of</strong> Schultze), but this, as shown by<br />

Gaupp, is a ventral process <strong>of</strong> Basibranchiale i, <strong>and</strong> so not homologous with<br />

<strong>the</strong> Urobranchiale <strong>of</strong> Urodela.)<br />

Transversi ventrales. Transversus ventralis i is present <strong>in</strong> larvae <strong>of</strong> Ichthyophis<br />

<strong>and</strong> Siph<strong>on</strong>ops, but not <strong>in</strong> Anuran or Urodelan larvae. Transversus<br />

ventralis ii is not found <strong>in</strong> any Amphibia. Transversus ventralis iii is present<br />

<strong>in</strong> Necturus, <strong>and</strong> probably <strong>in</strong> Proteus, but not <strong>in</strong> o<strong>the</strong>r Amphibia. Transversus<br />

ventralis iv is present <strong>in</strong> larvae <strong>of</strong> Anura, Gymnophi<strong>on</strong>a, <strong>and</strong> Urodela with<br />

four br<strong>on</strong>chial bars. In Rana <strong>the</strong> muscle is rudimentary <strong>and</strong> so<strong>on</strong> disappears.<br />

In Ichthyophis it is attached to <strong>the</strong> fourth br<strong>on</strong>chial bar, <strong>and</strong>, <strong>on</strong> <strong>the</strong> fusi<strong>on</strong><br />

<strong>of</strong> this with <strong>the</strong> third, to <strong>the</strong> fused bar, by two heads <strong>in</strong> Ichthyophis, by <strong>on</strong>e<br />

<strong>in</strong> Siph<strong>on</strong>ops.<br />

In Menopoma, at <strong>the</strong> stage when <strong>the</strong> sixth gill-cleft reaches <strong>the</strong> ectoderm,<br />

<strong>the</strong> ventral end <strong>of</strong> <strong>the</strong> fourth br<strong>on</strong>chial muscle-plate (attached to <strong>the</strong> pericardial<br />

wall) <strong>and</strong> <strong>the</strong> primordium <strong>of</strong> <strong>the</strong> fourth br<strong>on</strong>chial bar are <strong>in</strong> fr<strong>on</strong>t <strong>of</strong>

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