Developmental Biology 7th ed by Gilbert, Scott F

Book Preface

The Questions of Developmental Biology

According to Aristotle, the first embryologist known to history, science begins with wonder: “It is owing to wonder that people began to philosophize, and wonder remains the beginning of knowledge.” The development of an animal from an egg has been a source of wonder throughout history. The simple procedure of cracking open a chick egg on each successive day of its 3-week incubation provides a remarkable experience as a thin band of cells is seen to give rise to an entire bird. Aristotle performed this procedure and noted the formation of the major organs. Anyone can wonder at this remarkable yet commonplace phenomenon, but the scientist seeks to discover how development actually occurs. And rather than dissipating wonder, new understanding increases it.

Multicellular organisms do not spring forth fully formed. Rather, they arise by a relatively slow process of progressive change that we call development. In nearly all cases, the development of a multicellular organism begins with a single cell the fertilized egg, or zygote, which divides mitotically to produce all the cells of the body. The study of animal development has traditionally been called embryology, from that stage of an organism that exists between fertilization and birth. But development does not stop at birth, or even at adulthood. Most organisms never stop developing. Each day we replace more than a gram of skin cells (the older cells being sloughed off as we move), and our bone marrow sustains the development of millions of new red blood cells every minute of our lives. In addition, some animals can regenerate severed parts, and many species undero metamorphosis (such as the transformation of a tadpole into a frog, or a caterpillar into a butterfly). Therefore, in recent years it has become customary to speak of developmental biology as the discipline that studies embryonic and other developmental processes.

Development accomplishes two major objectives: it generates cellular diversity and order within each generation, and it ensures the continuity of life from one generation to the next. Thus, there are two fundamental questions in developmental biology: How does the fertilized egg give rise to the adult body, and how does that adult body produce yet another body? These two huge questions have been subdivided into six general questions scrutinized by developmental biologists:

The question of differentiation. A single cell, the fertilized egg, gives rise to hundreds of different cell types muscle cells, epidermal cells, neurons, lens cells, lymphocytes, blood cells, fat cells, and so on (Figure 1.1). This generation of cellular diversity is called differentiation. Since each cell of the body (with very few exceptions) contains the same set of genes, we need to understand how this same set of genetic instructions can produce different types of cells. How can the fertilized egg generate so many different cell types?

The question of morphogenesis. Our differentiated cells are not randomly distributed. Rather, they are organized into intricate tissues and organs. These organs are arranged in a given way: the fingers are always at the tips of our hands, never in the middle; the eyes are always in our heads, not in our toes or gut. This creation of ordered form is called morphogenesis. How can the cells form such ordered structures?

were to undergo just one more cell division, we would be considered horribly malformed. If each cell in our arms underwent just one more round of cell division, we could tie our shoelaces without bending over. Our arms are generally the same size on both sides of the body. How is cell division so tightly regulated?

The question of reproduction. The sperm and egg are very specialized cells. Only they can transmit the instructions for making an organism from one generation to the next. How are these cells set apart to form the next generation, and what are the instructions in the nucleus and cytoplasm that allow them to function this way?

The question of evolution. Evolution involves inherited changes in development. When we say that today’s one-toed horse had a five-toed ancestor, we are saying that changes in the development of cartilage and muscles occurred over many generations in the embryos of the horse’s ancestors. How do changes in development create new body forms? Which heritable changes are possible, given the constraints imposed by the necessity of the organism to survive as it develops?

The question of environmental integration. The development of many organisms is influenced by cues from the environment. Certain butterflies, for instance, inherit the ability to produce different wing colors based on the temperature or the amount of daylight experienced by the caterpillar before it undergoes metamorphosis. How is the development of an organism integrated into the larger context of its habitat?