fertilization.

Early development

Embryo formation

Cleavage

Since the goal of development is the production of a multicellular organism, many cells must be produced from the single-celled zygote. This task is accomplished by cleavage, a series of consecutive cell divisions. Cells produced during cleavage are called blastomeres. The divisions are mitotic—i.e., each chromosome in the nucleus splits into two daughter chromosomes, so that the two daughter blastomeres retain the diploid number of chromosomes. During cleavage, almost no growth occurs between consecutive divisions, and the total volume of living matter does not change substantially; as a consequence, the size of the cells is reduced by almost half at each division. At the beginning of cleavage, cell divisions tend to occur at the same time in all blastomeres, and the number of cells is doubled at each division. As cleavage progresses, the cells no longer divide at the same time.

Cleavage in most animals follows an orderly pattern, with the first division being in the plane of the main axis of the egg. This cleavage plane is arbitrarily called vertical, on the assumption that the main axis of the egg is vertical. The second cleavage plane is again vertical but at right angles to the first, giving rise to four equal cells arranged around the main axis of the egg. The third cleavage plane is at right angles to both the first and second cleavage planes and is horizontal, or equatorial. Subsequent divisions may alternate between vertical and horizontal cleavage planes, but later cleavage divisions become randomly oriented. This pattern is typical of many animal groups; however, more complicated patterns of cleavage are found in such animals as annelids, mollusks, and nematodes.

As the amount of yolk in the egg increases, it influences cleavage by hindering the cytoplasmic movements involved in mitosis. If there is only little yolk (oligolecithal eggs), the yolk granules follow the movements of the cytoplasm and are distributed in the resulting blastomeres. But if the amount of yolk is larger (megalecithal eggs), cleavages occur nearer the animal pole, where there is less yolk; as a result, the blastomeres nearer the animal pole are smaller than those nearer the vegetal pole. The presence of yolk masses may retard the onset of cleavage in a part of the egg or even suppress it altogether; in this case cleavage is partial, or meroblastic. Only a part of the egg material then is subdivided into cells, the rest remaining as a mass that serves as nourishment for the developing embryo.

Cleavage is complete, or holoblastic, in many invertebrates including coelenterates, annelids, echinoderms, tunicates, and cephalochordates. The blastomeres may be either about equal or only slightly different in size. Cleavage in amphibians is holoblastic, but the size of the blastomeres is very uneven. Blastomeres are smallest at the animal pole and largest (and yolky) at the vegetal pole. Somewhat similar conditions prevail in many mollusks. In most fishes, birds, reptiles, and egg-laying mammals (monotremes), cleavage is discoidal—i.e., restricted to a disk of cytoplasm at the animal pole of the egg, most of the yolky egg material remaining uncleaved. Cleavage in insects and many other arthropods is superficial—i.e., the entire surface layer of egg cytoplasm subdivides into cells, and the egg contains a central mass of uncleaved yolk. The conditions of cleavage in placental mammals, including man, are peculiar.

During cleavage, development involves only an increase of cell numbers; the shape of the embryo does not change, and chemical transformations within the embryo are restricted to those necessary for cell division. Chemical and structural transformations are concerned with accumulating chromosomal material in the nuclei of the blastomeres. Before each division the chromosomes carrying the genes double in number; this means that the chromosomal material, deoxyribonucleic acid (DNA), has to be synthesized. This synthesis proceeds possibly at the expense of cytoplasmic ribonucleic acid (RNA) but certainly also from simpler organic compounds. A certain amount of protein synthesis is also necessary for cleavage to proceed: if developing eggs are treated with puromycin, a substance which is known to suppress protein synthesis, cleavage stops immediately. The proteins concerned have not yet been identified. No proteins are synthesized, however, that would foreshadow the future differentiation of parts of the embryo. It is believed that the genes in the chromosomes remain largely inactive during cleavage. The rhythm (speed) of cleavage is wholly dependent on the cytoplasm of the egg.

Although the shape and volume of the embryo do not change during cleavage, one important change in gross organization does take place. As the blastomeres are produced, they move outward, leaving a centrally located fluid-filled cavity. In cases of holoblastic cleavage, the blastomeres become arranged in a layer from one to several cells thick surrounding the cavity. The embryo at this stage may be likened to a hollow ball and is known as a blastula. The outer layer of cells is called the blastoderm, and the fluid-filled cavity the blastocoel. In discoidal cleavage the cells, which do not surround the whole embryo, lie only on the animal pole; nevertheless, a blastocoel may be formed by a crevice appearing between the blastomeres and the mass of yolk. The blastomeres then may be arranged as a saucer-shaped blastodisk covering the blastocoel.

The formation of the blastula signifies the end of the period of cleavage. The next stage of development is concerned not with an increase in cell number, though cell divisions continue at a slower pace, but with rearrangement of the available cell masses to conform with the gross features of the future animal.

Gastrulation

The embryo in the blastula stage must go through profound transformations before it can approach adult organization. An adult multicellular animal typically possesses a concentric arrangement of tissues of the body; this feature is common to all animal groups above the level of the sponges. Adult tissues are derived from three embryonic cell layers called germinal layers: the outer layer is the ectoderm, the middle layer is the mesoderm, and the innermost layer is the endoderm (entoderm). The ectoderm gives rise to the skin covering, to the nervous system, and to the sense organs. The mesoderm produces the muscles, excretory organs, circulatory organs, sex organs (gonads), and internal skeleton. The endoderm lines the alimentary canal and gives rise to the organs associated with digestion and, in chordates, with breathing.

The blastula, which consists of only one cell layer, undergoes a dramatic reshuffling of blastomeres preparatory to the development of the various organ systems of the animal’s body. This is achieved by the process of gastrulation, which is essentially a shifting or moving of the cell material of the embryo so that the three germinal layers are aligned in their correct positions.

The rearrangement of the blastula to form the germinal layers is seen clearly in certain marine animals with oligolecithal eggs. The hollow blastula consists of a simple epithelial layer (the blastoderm), the transformation of which can be likened to the pushing in of one side of a rubber ball. As a result of such inpushing (or invagination), the spherical embryo is converted into a double-walled cup, the opening of which represents the position of the former vegetal pole. The involuted part of the blastoderm, lining the inside of the double-walled cup, gives rise to the endoderm and mesoderm, and the blastomeres remaining on the exterior become the ectoderm. As a consequence of the infolding at the vegetal pole, the blastocoel is reduced or obliterated, and a new cavity is created, the primitive gut cavity, or archenteron, which eventually gives rise to the hollow core (lumen) of the alimentary canal. At this stage the embryo has a primitive gut with an opening to the exterior and is known as a gastrula. The opening of the gastrula is the blastopore, or primitive mouth; both terms are somewhat misleading. It would seem that the term blastopore should be applied more appropriately to an opening in a blastula, in which, of course, no opening exists. As to the term primitive mouth, it must be pointed out that the blastopore does not always give rise to the adult mouth. In certain animal groups it becomes the anus, and a mouth forms as a completely new opening.

In some coelenterates, cells at the vegetal pole do not form an invaginating pocket, but individual cells slide inward, losing connection with other cells of the blastoderm. Eventually these cells fill the blastocoel and form a compact mass of endoderm. The cavity (archenteron) within this mass and the opening (blastopore) to the exterior are then produced secondarily by the separation of these cells.