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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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CHAPTER 21 END-OF-CHAPTER PROBLEMS

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fundamental role in learning and memory. Clearly, for the nervous system as for

other parts of the body, developmental processes do not end at birth, as we discuss

in the next chapter.

Summary

The development of the nervous system proceeds in four phases. First, neurons and

glial cells are generated from dividing neural progenitor cells. Then, the newborn

neurons send out axons and dendrites toward their targets. Next, they make synaptic

connections with appropriate target cells so that communication can begin.

Finally, excessive neurons are eliminated by normal neuronal cell death, after

which the system of synaptic connections is refined and remodeled according to the

pattern of electrical and synaptic activity in the neural network.

Neurons born at different times and places are specialized to express different

sets of genes, and they have a cell memory that plays a major role in determining

the connections they will form. Their specialization depends not only on spatial

patterning by morphogens but also on intrinsic developmental programs that

unfold as the neural progenitors proliferate. Axons and dendrites grow out from

the neurons by means of growth cones, which follow specific pathways delineated

by attractive and repellant signals along the way, including cell-surface and extracellular

matrix molecules and soluble signal proteins to which growth cones from

different classes of neurons respond differently. In many parts of the nervous system,

neural maps are set up—orderly projections of one array of neurons onto another.

In the retinotopic system, the map is based on the matching of complementary systems

of position-specific cell-surface markers—ephrins and Eph receptors—possessed

by the two sets of cells. Other cell-surface molecules such as DSCAM proteins

in Drosophila and protocadherins in vertebrates mediate self-avoidance between

the branches arising from a single neuron, helping the cell spread out its processes.

The formation of synapses involves back-and-forth signaling between target

cells and the growth cone. After the growth cones have reached their targets and

initial connections have formed, individual synapses are eliminated in some places

and reinforced in others by mechanisms that depend on synaptic and electrical

activity. These mechanisms adjust the architecture of the neural network according

to the way in which it is used.

What we don't know

• What regulates the pace of

development? Why does a mouse

embryo develop faster than a human

embryo, for example?

• What are the mechanisms that

allow cell memory to be stored during

development, explaining how each

cell’s history determines its future

behavior?

• How do signals move through

tissues? What are the roles of the

extracellular matrix and of elongated

cell projections?

• How does a cell know exactly where

it is in a multicellular organism? How

does it know that its neighbors are the

correct ones and that, if not, it should

move or kill itself?

• How do cells respond to tiny

gradients of molecules in their

environment, as required for knowing

their positions? How are morphogen

gradients reliably interpreted?

• What are the genetic changes that

allow the repurposing of existing body

parts during evolution? For example,

how did bat wings evolve from arms?

• How do cells use genetic

instructions to form the shape of

something as complex as the human

nose?

Problems

Which statements are true? Explain why or why not.

21–1 In the early cleavage stages, when the embryo

cannot yet feed, the developmental program is driven and

controlled entirely by the material deposited in the egg by

the mother.

21–2 Because of the many later developmental transformations

that produce the elaborately structured organs,

the body plan set up during gastrulation bears little resemblance

to the body plan in the adult.

21–3 As development progresses, individual cells

become more and more restricted in the range of cell types

they can give rise to.

21–4 At different stages of embryonic development,

the same signals are used over and over again by different

cells, but with different biological outcomes.

21–5 Changes in the coding regions of genes involved in

development are primarily responsible for the differences

between species.

21–6 The cell cycle is the ticking clock that sets the

tempo of developmental processes, with maturational

changes in gene expression being dependent on cell-cycle

progression.

Discuss the following problems.

21–7 Name the four processes that are fundamental to

animal development, and describe each of them in a single

sentence.

21–8 What are the three germ layers formed during gastrulation,

and what are the principal structures each gives

rise to in the adult?

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