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The Questions of Developmental Biology

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embryos can produce these cells, but they appear unable to organize them into functional tissues<br />

and organs. Tissues <strong>of</strong> the digestive tract can be found, but they are not connected; muscle cells<br />

are scattered around the lobeless larva, but are not organized into a functional muscle tissue.<br />

Thus, the developmental functions <strong>of</strong> the polar lobe may be very complex.<br />

Fate map <strong>of</strong> Ilyanassa obsoleta<br />

As detailed above, fate maps <strong>of</strong> the snail Ilyanassa have been made means <strong>of</strong> ablation<br />

experiments. More recently, Joanne Render (1997) has constructed a more detailed fate map by<br />

injecting specific micromeres with beads containing the fluorescent dye Lucifer Yellow.<br />

<strong>The</strong> fluorescence is<br />

maintained over the period <strong>of</strong><br />

embryogenesis and can be seen in<br />

the larval tissue derived from the<br />

injected cells. Figure 8.35 shows the<br />

new fate map <strong>of</strong> Ilyanassa obsoleta.<br />

<strong>The</strong> second quartet micromeres (2ad)<br />

generally contribute to the shellforming<br />

mantle, the velum, the<br />

mouth, and the heart. <strong>The</strong> third<br />

quartet micromeres (3a-d) generate<br />

large regions <strong>of</strong> the foot, velum,<br />

esophagus, and heart. <strong>The</strong> 4d cell,<br />

the mesentoblast, contributes to the<br />

larval kidney, heart, retractor<br />

muscles, and intestine.<br />

Gastrulation in Snails<br />

<strong>The</strong> snail stereoblastula is<br />

relatively small, and its cell fates<br />

have already been determined by<br />

the D series <strong>of</strong> macromeres.<br />

Gastrulation is accomplished<br />

primarily by epiboly, wherein the<br />

micromeres at the animal cap<br />

multiply and "overgrow" the vegetal<br />

macromeres. Eventually, the<br />

micromeres will cover the entire<br />

embryo, leaving a small slit at the<br />

vegetal pole (Figure 8.36; Collier<br />

1997).

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