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The Questions of Developmental Biology

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This occurred at precisely the time that epiboly would have taken place during normal<br />

gastrulation. Thus, epiboly may be (at least in some respects) independent <strong>of</strong> the cells that form<br />

the migrating region.<br />

Axis Formation<br />

Some <strong>of</strong> the most important phenomena<br />

in development concern the formation <strong>of</strong><br />

embryonic axes (Figure 8.7). Embryos<br />

must develop three very important axes<br />

that are the foundations <strong>of</strong> the body: the<br />

anterior-posterior axis, the dorsalventral<br />

axis, and the right-left axis.<br />

<strong>The</strong> anterior-posterior (or<br />

anteroposterior) axis is the line<br />

extending from head to tail (or mouth to<br />

anus in those organisms that lack heads<br />

and tails).<br />

<strong>The</strong> dorsal-ventral (dorsoventral) axis is the<br />

line extending from back (dorsum) to belly (ventrum).<br />

For instance, in vertebrates, the neural tube is a dorsal<br />

structure. In insects, the neural cord is a ventral<br />

structure. <strong>The</strong> right-left axis is a line between the two<br />

lateral sides <strong>of</strong> the body. Although we may look<br />

symmetrical, recall that in most <strong>of</strong> us, the heart and liver<br />

are in the left half <strong>of</strong> the body only. Somehow, the<br />

embryo knows that some organs go on one side and<br />

other organs go on the other.<br />

In this chapter, we will look at how four selected invertebrates a sea urchin, a tunicate,<br />

a snail, and a nematode undergo cleavage, gastrulation, axis specification, and cell fate<br />

determination. <strong>The</strong>se four invertebrates were chosen because they have been important model<br />

systems for developmental biologists. In other words, they can be studied easily in the laboratory,<br />

and they have special properties that allow their mechanisms <strong>of</strong> development to be readily<br />

observed. <strong>The</strong>y also represent a wide variety <strong>of</strong> cleavage types, patterns <strong>of</strong> gastrulation, and ways<br />

<strong>of</strong> specifying axes and cell fates.*<br />

*However, model systems by their very ability to develop in the laboratory preclude our asking certain questions<br />

concerning the relationship <strong>of</strong> development to its habitat. <strong>The</strong>se questions will be addressed in Chapter 21<br />

<strong>The</strong> Early Development <strong>of</strong> Sea Urchins<br />

Cleavage in Sea Urchins<br />

Sea urchins exhibit radial holoblastic cleavage. <strong>The</strong> first and second cleavages are both<br />

meridional and are perpendicular to each other. That is to say, the cleavage furrows pass through<br />

the animal and vegetal poles. <strong>The</strong> third cleavage is equatorial, perpendicular to the first two<br />

cleavage planes, and separates the animal and vegetal hemispheres from one another (Figures 8.8<br />

and 8.9). <strong>The</strong> fourth cleavage, however, is very different from the first three.

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