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The Questions of Developmental Biology

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<strong>The</strong> surfactant enables the alveolar cells to touch one another without sticking together.<br />

Thus, infants born prematurely <strong>of</strong>ten have difficulty breathing and have to be placed on<br />

respirators until their surfactant-producing cells mature.<br />

As in the digestive tube, the regional specificity <strong>of</strong> the<br />

mesenchyme determines the differentiation <strong>of</strong> the developing<br />

respiratory tube. In the developing mammal, the respiratory<br />

epithelium responds in two distinct fashions. In the region <strong>of</strong> the<br />

neck, it grows straight, forming the trachea. After entering the<br />

thorax, it branches, forming the two bronchi and then the lungs. <strong>The</strong><br />

respiratory epithelium can be isolated soon after it has split into two<br />

bronchi, and the two sides can be treated differently. Figure 15.32<br />

shows the result <strong>of</strong> such an experiment. <strong>The</strong> right bronchial<br />

epithelium retained its lung mesenchyme, whereas the left bronchus<br />

was surrounded with tracheal mesenchyme (Wessells 1970). <strong>The</strong> right bronchus proliferated and<br />

branched under the influence <strong>of</strong> the lung mesenchyme, whereas the left bronchus continued to<br />

grow in an unbranched manner. Moreover, the differentiation <strong>of</strong> the respiratory epithelia into<br />

trachea or lung cells depends on the mesenchyme it encounters (Shannon et al. 1998). Thus, the<br />

respiratory epithelium is extremely malleable and can differentiate according to its mesenchymal<br />

instructions.<br />

<strong>The</strong> Extraembryonic Membranes<br />

In reptiles, birds, and mammals, embryonic development has taken a new evolutionary<br />

direction. Reptiles evolved a mechanism for laying eggs on dry land, thus freeing them to explore<br />

niches that were not close to water. To accomplish this, the embryo produces four sets <strong>of</strong><br />

extraembryonic membranes to mediate between it and the environment (see Chapter 11). Even<br />

though most mammals have evolved placentas instead <strong>of</strong> shells, the basic pattern <strong>of</strong><br />

extraembryonic membranes remains the same. In developing reptiles, birds, and mammals, there<br />

initially is no distinction between embryonic and extraembryonic domains. However, as the body<br />

<strong>of</strong> the embryo takes shape, the epithelia at the border between the embryo and the extraembryonic<br />

domain divide unequally to create body folds that isolate the embryo from the yolk and delineate<br />

which areas are to be embryonic and which extraembryonic (Miller et al. 1994, 1999). <strong>The</strong>se<br />

membranous folds are formed by the extension <strong>of</strong> ectodermal and endodermal epithelium<br />

underlain with lateral plate mesoderm. <strong>The</strong> combination <strong>of</strong> ectoderm and mesoderm, <strong>of</strong>ten<br />

referred to as the somatopleure, forms the amnion and chorion; the combination <strong>of</strong> endoderm<br />

and mesoderm the splanchnopleure forms the yolk sac and allantois. <strong>The</strong> endodermal or<br />

ectodermal tissue supplies functioning epithelial cells, and the mesoderm generates the essential<br />

blood supply to and from this epithelium. <strong>The</strong> formation <strong>of</strong> these folds can be followed in Figure<br />

15.33.<br />

<strong>The</strong> amnion and chorion<br />

<strong>The</strong> first problem <strong>of</strong> a land-dwelling egg is desiccation. Embryonic cells would quickly dry out if<br />

they were not in an aqueous environment. This environment is supplied by the amnion. <strong>The</strong> cells<br />

<strong>of</strong> this membrane secrete amnionic fluid; thus, embryogenesis still occurs in water. This<br />

evolutionary adaptation is so significant and characteristic that reptiles, birds, and mammals are<br />

grouped together as the amniote vertebrates, or amniotes.

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