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The Questions of Developmental Biology

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<strong>The</strong> <strong>Developmental</strong> Mechanics <strong>of</strong> Cell Specification<br />

An embryo's environment may be a tide pool, a pond, or a uterus. As we saw above, the<br />

embryo interacts with its environment, and its developmental trajectory can be guided by<br />

information from its surroundings. On a smaller scale, the environment <strong>of</strong> an embryonic cell<br />

consists <strong>of</strong> the surrounding tissues within the embryo, and the fate <strong>of</strong> that cell (for instance,<br />

whether it becomes part <strong>of</strong> the skin or part <strong>of</strong> the lens) <strong>of</strong>ten depends upon its interactions with<br />

other components <strong>of</strong> its immediate "ecosystem."<br />

Thus, a second research program <strong>of</strong> experimental embryology studies how interactions<br />

between embryonic cells generate the embryo. <strong>The</strong> development <strong>of</strong> specialized cell types is called<br />

differentiation (Table 3.2). <strong>The</strong>se overt changes in cellular biochemistry and function are<br />

preceded by a process involving the commitment <strong>of</strong> the cell to a certain fate. At this point, even<br />

though the cell or tissue does not appear phenotypically different from its uncommitted state, its<br />

developmental fate has become restricted.<br />

Table 3.2. Some differentiated cell types and their major produts<br />

Cell type Differentiated cell product Specialized function <strong>of</strong> cell<br />

Keratinocyte (epidermal Keratin<br />

Protection against abrasion, desiccation<br />

cell)<br />

Erythrocyte (red blood Hemoglobin<br />

Transport <strong>of</strong> oxygen<br />

cell)<br />

Lens cell Crystallins Transmission <strong>of</strong> light<br />

B lymphocyte Immunoglobulins Antibody synthesis<br />

T lymphocyte Cell-surface antigens (lymphokines) Destruction <strong>of</strong> foreign cells; regulation <strong>of</strong> immune<br />

response<br />

Melanocyte Melanin Pigment production<br />

Pancreatic islet cells Insulin Regulation <strong>of</strong> carbohydrate metabolism<br />

Leydig cell ( ) Testosterone Male sexual characteristics<br />

Chondrocyte (cartilage Chondroitin sulfate; type II collagen Tendons and ligaments<br />

cell)<br />

Osteoblast (bone-forming Bone matrix<br />

Skeletal support<br />

cell)<br />

Myocyte (muscle cell) Muscle actin and myosin Contraction<br />

Hepatocyte (liver cell) Serum albumin; numerous enzymes Production <strong>of</strong> serum proteins and numerous<br />

enzymatic functions<br />

Neurons Neurotransmitters (acetylcholine Transmission <strong>of</strong> electrical impulses<br />

epinephrine, etc.)<br />

Tubule cell ( ) <strong>of</strong> hen Ovalbumin<br />

Egg white proteins for nutrition and protection <strong>of</strong><br />

oviduct<br />

embryo<br />

Follicle cell ( ) <strong>of</strong> insect Chorion proteins<br />

Eggshell proteins for protection <strong>of</strong> embryo<br />

ovary<br />

<strong>The</strong> process <strong>of</strong> commitment can be divided into two stages (Harrison 1933; Slack 1991).<br />

<strong>The</strong> first stage is a labile phase called specification. <strong>The</strong> fate <strong>of</strong> a cell or a tissue is said to be<br />

specified when it is capable <strong>of</strong> differentiating autonomously when placed in a neutral<br />

environment such as a petri dish or test tube. (<strong>The</strong> environment is neutral with respect to the<br />

developmental pathway.) At this stage, the commitment is still capable <strong>of</strong> being reversed. <strong>The</strong><br />

second stage <strong>of</strong> commitment is determination. A cell or tissue is said to be determined when it is

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