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Cleavage and gastrulation in the shrimp Penaeus (Litopenaeus ...

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Fig. 1. Cell l<strong>in</strong>eage of <strong>Penaeus</strong> vannamei, show<strong>in</strong>g <strong>the</strong> relative tim<strong>in</strong>g of cleavage divisions <strong>in</strong> m<strong>in</strong> ps at 27 8C (A) <strong>and</strong> naupliar mesoderm <strong>and</strong> mesendoderm<br />

detail (B–E). (A) The overall l<strong>in</strong>eage is color coded as to germ layer, with ectoderm <strong>in</strong> blue, mesoderm <strong>in</strong> red, endoderm <strong>in</strong> yellow, <strong>and</strong> germ l<strong>in</strong>e <strong>in</strong> purple. The<br />

nomenclature refers to <strong>the</strong> most commonly observed cleavage pattern, where DIIrZX, <strong>the</strong> progenitor of <strong>the</strong> two mesendoderm cells. The AB blastomere<br />

conta<strong>in</strong>s <strong>the</strong> animal pole, while <strong>the</strong> CD blastomere conta<strong>in</strong>s <strong>the</strong> vegetal pole. A <strong>and</strong> C make cross-furrow contact, as do B <strong>and</strong> D. The subscript I refers to <strong>the</strong><br />

cell adjacent to <strong>the</strong> cross-furrow (e.g. A I), while <strong>the</strong> subscript II refers to <strong>the</strong> cell away from <strong>the</strong> cross furrow (e.g. A II). Additional designations are based on relative<br />

axial positions, ei<strong>the</strong>r anterior–posterior (a, p), dorsal–ventral (d, v) or left–right (l, r). L<strong>in</strong>eage of <strong>the</strong> ectoderm is shown through 7 cell divisions. The l<strong>in</strong>eage <strong>and</strong><br />

nomenclature of <strong>the</strong> 9 crown cells K1–K9 are shown through two additional divisions. The derivatives of <strong>the</strong> mesendoderm are: <strong>the</strong> endodermal yolk cells,<br />

primordial endoderm cell E, mesoteloblast M, <strong>and</strong> germ cell G. (B) 122-cell stage, vegetal view show<strong>in</strong>g replicated crown cells. (C) Dorsal (X d) <strong>and</strong> ventral (X v)<br />

mesendoblasts, with <strong>the</strong> rest of <strong>the</strong> embryo omitted for clarity. (D) Result of first (anterior–posterior) mesendoderm division, posterior view. (E) Result of second<br />

mesendoderm division, posterior view. Endodermal yolk cells (Xdpd, Xdal, Xdar, Xval, Xvar), primordial endoderm cell (E), primordial germ cell (G), <strong>and</strong><br />

primordial mesoteloblast (M) are labeled. (F) Result of third mesendoderm division, posterior view. (G) Result of fourth mesendoderm division, posterior view.<br />

<strong>and</strong> BD b<strong>and</strong>s formed two rows of four cells with sp<strong>in</strong>dles<br />

oriented end-to-end, result<strong>in</strong>g <strong>in</strong> a 32-cell stage embryo<br />

(Fig. 2(M)–(P), Fig. 3(A), (D) <strong>and</strong> (G)). The AC <strong>and</strong> BD<br />

b<strong>and</strong>s divided dorsal–ventrally. AIIa formed AIIad <strong>and</strong><br />

P.L. Hertzler / Arthropod Structure & Development 34 (2005) 455–469<br />

AIIav, AIIp formed AIIpd <strong>and</strong> AIIpv, AIa formed AIad <strong>and</strong><br />

AIav, <strong>and</strong> AIp formed AIpd <strong>and</strong> AIpv (Fig. 3(B), (C), (E) <strong>and</strong><br />

(F)). CIIa formed CIIad <strong>and</strong> CIIav, CIIp formed CIIpd<br />

<strong>and</strong> CIIpv, CIa formed CIad <strong>and</strong> CIav, <strong>and</strong> CIp formed CIpd

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