Activity 47.1 What common events occur in the early development of animals? 1. What key events occur at each stage of development?

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1 Notes to Instructors Chapter 47 Animal Development What is the focus of this activity? Chapter 21 provided a review of how genes act to control development. Chapter 47 reviews some of the major morphological changes that occur in the development of many animals. Similarities in early development are often used as evidence for evolutionary relationships among organisms. What is this particular activity designed to do? Activity 47.1 What common events occur in the early development of animals? This activity provides a brief overview of major events that occur in the early development of vertebrates and many other animals. Answers Activity 47.1 What common events occur in the early development of animals? The early stages in the development of all vertebrates (and many other animals) include zygote formation, cleavage, blastula formation, gastrula formation, and organogenesis (for example, neurulation). Among the major aspects of development are cell adhesion/ recognition, cell growth (in number and/or size), cell induction, and cell determination. 1. What key events occur at each stage of development? Developmental stage What occurs during this stage? a. Cleavage In vertebrates, cell division is radial and indeterminate. The first mitotic cell divisions that follow fertilization occur rapidly and are not accompanied by cell growth. The What is the influence or effect on the subsequent development of the embryo? In many animals (mammals are an exception), the zygote has a definite polarity. As a result, the cleavage divisions create cells that differ in their cytoplasmic contents (e.g., cytoplasmic determinants: mrnas and proteins present in the egg cell). (Continued on next page) 310 Notes to Instructors

2 b. Blastula formation c. Gastrula formation chromosomes duplicate and are partitioned to daughter cells, but the cytoplasm is simply partitioned among the daughter cells. As a result, these divisions are called cleavage. As cleavage continues, the blastula forms. The blastula is a hollow ball of cells surrounding a fluid-filled cavity, or blastocoel. Blastula cells (near what will become the blastopore) begin to divide rapidly. These cells undergo changes in their shape, motility, and adhesion properties. Overall, there is a general migration of cells into the blastocoel to form the archenteron (primitive gut). Ultimately, gastrulation results in the formation of the germ layers: ectoderm, endoderm, and mesoderm. These differences can affect the subsequent development of the daughter cells. Cells on the outside of the blastula are in direct contact with the external environment. Those on the inside are in contact with the fluid-filled blastocoel. Those that lie between are only in contact with other cells. As a result, the cells experience differences in chemical and physical factors that can affect how each develops. These cellular movements and rearrangements produce new gradients in chemical signals and new types of cell-cell interactions between ectoderm and mesoderm and between mesoderm and endoderm. Extracellular glycoproteins, such as fibronectin, have been shown to play a role in directing the cell movements of gastrulation. By the late gastrula stage, the fates of many of the cells have been determined. In general, ectodermal cells develop into the nervous system and the epidermis. The endodermal cells become the digestive tract and associated organs (liver and pancreas).mesodermal cells give rise to the muscles, the circulatory system, the kidneys, and the dermis. In deuterostomes (echinoderms and chordates), the anus forms near the old blastopore and the mouth forms from a new opening (opposite the site of the blastopore). (Continued on next page) Activity

3 Developmental stage d. Organogenesis (for example, neural tube formation) What occurs during this stage? Organogenesis is basically the formation of the organs from the germ layers set up in the gastrula. The notochord and neural tube (nervous system) are among the first organ systems to develop in vertebrates. What is the influence or effect on the subsequent development of the embryo? The development of one structure is often triggered by or dependent on signals (chemical or physical) produced by other structures. For example, in early development, the cells of the dorsal lip of the blastopore appear to initiate the development of the neural tube and a number of other organs. 2. Many animal species share these similarities in early development. Yet, the stage at which the individual cells of the embryo lose their totipotency can vary considerably among these species. a. What does it mean to say that a cell is totipotent? A cell that is totipotent is undifferentiated and has the potential to differentiate into any of a number of different types of cell. b. What factors can affect the point at which a cell loses its totipotency that is, when its fate becomes determined? Among the factors that affect when a cell loses it totipotency are: the distribution of cytoplasmic determinants in the cells, the presence of organizing centers or regions that appear to trigger the development of surrounding cells, an unequal distribution of chemical signals, and specific cell-cell interactions encountered by the cell. Each of these factors appears to have some effect on which genes are activated in the cells during development and, as a result, how the cells differentiate. 312 Activity 47.1

4 47.1 Test Your Understanding 1 and 2. Choose the graph that best fits each situation. A B C D E F 1. Which graph best describes the change in size of individual cells of a vertebrate embryo from the of zygote formation to the end of cleavage? Explain your answer. D During cleavage cells divide rapidly, but there is little or no intervening cell growth between mitoses. As a result, the cells become smaller and smaller with each division. 2. Which graph best describes the change in the number of cells in the embryo from fertilization to gastrulation? Explain your answer. B The overall number of cells in the embryo increases continuously from fertilization to gastrulation. 3 to 6. While the mechanics of gastrulation may differ among various organisms, the overall objectives and problems are the same. Which of the following are accomplished by the end of gastrulation? T/F 3. formation of the three germ layers ectoderm, mesoderm, and endoderm T/F 4. establishment of the embryonic axis (anterior to posterior) T/F 5. determination of the fates of the individual cells of the gastrula T/F 6. formation of the neural tube False This occurs as part of organogenesis Activity

5 7. In a now classic experiment, Spemann and Mangold took cells from the dorsal lip of the blastopore from one frog embryo (the donor) and transplanted them to an area opposite the dorsal lip in another frog embryo (the host). As a result, the host embryo developed two heads, one at the site of the new or transplanted dorsal lip and the other at the site of the host s original dorsal lip. A thorough examination indicated that the second head was formed from host cells. Which of the following developmental cues or mechanisms was most likely the trigger (or cause) for the generation of the second head. Explain your answer. a. cytoplasmic determinants that are unequally distributed in the host embryo b. gradients set up in the egg that gave rise to the host embryo c. hox genes (or homeobox genes) present in the embryo receiving the transplant d. cell-cell signalling between the transplanted dorsal lip and the host embryo e. all of the above d If the donor cells had not been transplanted to the host, the host would have developed normally. As a result, the altered development must have been triggered by some chemical or cell-cell signal from the transplanted or donor tissue. 314 Activity 47.1

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