153 Embryological Development

Most animal species undergo a separation of tissues into germ layers during embryonic development. Recall that these germ layers are formed during gastrulation, and that they are predetermined to develop into the animal’s specialized tissues and organs. Animals develop either two or three embryonic germs layers (Figure 1). The animals that display radial symmetry develop two germ layers, an inner layer (endoderm) and an outer layer (ectoderm). These animals are called diploblasts. Diploblasts have a non-living layer between the endoderm and ectoderm. More complex animals (those with bilateral symmetry) develop three tissue layers: an inner layer (endoderm), an outer layer (ectoderm), and a middle layer (mesoderm). Animals with three tissue layers are called triploblasts.

Each of the three germ layers is programmed to give rise to particular body tissues and organs. The endoderm gives rise to the lining of the digestive tract (including the stomach, intestines, liver, and pancreas), as well as to the lining of the trachea, bronchi, and lungs of the respiratory tract, along with a few other structures. The ectoderm develops into the outer epithelial covering of the body surface, the central nervous system, and a few other structures. The mesoderm is the third germ layer; it forms between the endoderm and ectoderm in triploblasts. This germ layer gives rise to all muscle tissues (including the cardiac tissues and muscles of the intestines), connective tissues such as the skeleton and blood cells, and most other visceral organs such as the kidneys and the spleen.

Presence or Absence of a Coelom

Further subdivision of animals with three germ layers (triploblasts) results in the separation of animals that may develop an internal body cavity derived from mesoderm, called a coelom, and those that do not. This epithelial cell-lined coelomic cavity represents a space, usually filled with fluid, which lies between the visceral organs and the body wall. It houses many organs such as the digestive system, kidneys, reproductive organs, and heart, and contains the circulatory system. In some animals, such as mammals, the part of the coelom called the pleural cavity provides space for the lungs to expand during breathing. The evolution of the coelom is associated with many functional advantages. Primarily, the coelom provides cushioning and shock absorption for the major organ systems. Organs housed within the coelom can grow and move freely, which promotes optimal organ development and placement. The coelom also provides space for the diffusion of gases and nutrients, as well as body flexibility, promoting improved animal motility.

Triploblasts that do not develop a coelom are called acoelomates, and their mesoderm region is completely filled with tissue, although they do still have a gut cavity. Examples of acoelomates include animals in the phylum Platyhelminthes, also known as flatworms. Animals with a true coelom are called eucoelomates (or coelomates) (Figure 2). A true coelom arises entirely within the mesoderm germ layer and is lined by an epithelial membrane. This membrane also lines the organs within the coelom, connecting and holding them in position while allowing them some free motion. Annelids, mollusks, arthropods, echinoderms, and chordates are all eucoelomates. A third group of triploblasts has a slightly different coelom derived partly from mesoderm and partly from endoderm, which is found between the two layers. Although still functional, these are considered false coeloms, and those animals are called pseudocoelomates. The phylum Nematoda (roundworms) is an example of a pseudocoelomate. True coelomates can be further characterized based on certain features of their early embryological development.

Embryonic Development of the Mouth

Bilaterally symmetrical, tribloblastic eucoelomates can be further divided into two groups based on differences in their early embryonic development. Protostomes include arthropods, mollusks, and annelids. Deuterostomes include more complex animals such as chordates but also some simple animals such as echinoderms. These two groups are separated based on which opening of the digestive cavity develops first: mouth or anus. The word protostome comes from the Greek word meaning “mouth first,” and deuterostome originates from the word meaning “mouth second” (in this case, the anus develops first). The mouth or anus develops from a structure called the blastopore (Figure 3).

The blastopore is the indentation formed during the initial stages of gastrulation. In later stages, a second opening forms, and these two openings will eventually give rise to the mouth and anus (Figure 3). It has long been believed that the blastopore develops into the mouth of protostomes, with the second opening developing into the anus; the opposite is true for deuterostomes. Recent evidence has challenged this view of the development of the blastopore of protostomes, however, and the theory remains under debate.

Another distinction between protostomes and deuterostomes is the method of coelom formation, beginning from the gastrula stage. The coelom of most protostomes is formed through a process called schizocoely, meaning that during development, a solid mass of the mesoderm splits apart and forms the hollow opening of the coelom. Deuterostomes differ in that their coelom forms through a process called enterocoely. Here, the mesoderm develops as pouches that are pinched off from the endoderm tissue. These pouches eventually fuse to form the mesoderm, which then gives rise to the coelom.

The earliest distinction between protostomes and deuterostomes is the type of cleavage undergone by the zygote. Protostomes undergo spiral cleavage, meaning that the cells of one pole of the embryo are rotated, and thus misaligned, with respect to the cells of the opposite pole. This is due to the oblique angle of the cleavage. Deuterostomes undergo radial cleavage, where the cleavage axes are either parallel or perpendicular to the polar axis, resulting in the alignment of the cells between the two poles.

There is a second distinction between the types of cleavage in protostomes and deuterostomes. In addition to spiral cleavage, protostomes also undergo determinate cleavage. This means that even at this early stage, the developmental fate of each embryonic cell is already determined. A cell does not have the ability to develop into any cell type. In contrast, deuterostomes undergo indeterminate cleavage, in which cells are not yet pre-determined at this early stage to develop into specific cell types. These cells are referred to as undifferentiated cells. This characteristic of deuterostomes is reflected in the existence of familiar embryonic stem cells, which have the ability to develop into any cell type until their fate is programmed at a later developmental stage.