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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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648 Chapter 12: Intracellular Compartments and Protein Sorting

Table 12–3 Some Typical Signal Sequences

Function of signal

sequence

Import into nucleus

Export from nucleus

Import into mitochondria

Example of signal sequence

-Pro-Pro-Lys-Lys-Lys-Arg-Lys-Val-

-Met-Glu-Glu-Leu-Ser-Gln-Ala-Leu-Ala-Ser-Ser-Phe-

+ H 3 N-Met-Leu-Ser-Leu-Arg-Gln-Ser-Ile-Arg-Phe-Phe-Lys-Pro-Ala-Thr-Arg-Thr-Leu-Cys-Ser-Ser-

Arg-Tyr-Leu-Leu-

Import into plastid

Import into peroxisomes -Ser-Lys-Leu-COO –

Import into ER

Return to ER -Lys-Asp-Glu-Leu-COO –

+ H 3 N-Met-Val-Ala-Met-Ala-Met-Ala-Ser-Leu-Gln-Ser-Ser-Met-Ser-Ser-Leu-Ser-Leu-Ser-Ser-Asn-

Ser-Phe-Leu-Gly-Gln-Pro-Leu-Ser-Pro-Ile-Thr-Leu-Ser-Pro-Phe-Leu-Gln-Gly-

+ H 3 N-Met-Met-Ser-Phe-Val-Ser-Leu-Leu-Leu-Val-Gly-Ile-Leu-Phe-Trp-Ala-Thr-Glu-Ala-Glu-Gln-

Leu-Thr-Lys-Cys-Glu-Val-Phe-Gln-

Some characteristic features of the different classes of signal sequences are highlighted in color. Where they are known to be important for

the function of the signal sequence, positively charged amino acids are shown in red and negatively charged amino acids are shown in green.

Similarly, important hydrophobic amino acids are shown in orange and important hydroxylated amino acids are shown in blue. + H 3 N indicates the

N-terminus of a protein; COO – indicates the C-terminus.

recognize classes of proteins rather than an individual protein species. They can

therefore be viewed as public transportation systems, dedicated to delivering

numerous different components to their correct location in the cell.

Most Organelles Cannot Be Constructed De Novo: They Require

Information in the Organelle Itself

When a cell reproduces by division, it has to duplicate its organelles, in addition to

its chromosomes. In general, cells do this by incorporating new molecules into the

existing organelles, thereby enlarging them; the enlarged organelles then divide

and are distributed to the two daughter cells. Thus, each daughter cell inherits a

complete set of specialized cell membranes from its mother. This inheritance is

essential because a cell could not make such membranes from scratch. If the ER

were completely removed from a cell, for example, how could the cell reconstruct

it? As we discuss later, the membrane proteins that define the ER and perform

many of its functions are themselves products of the ER. A new ER could not be

made without an existing ER or, at least, a membrane that specifically contains

the protein translocators required to import selected proteins into the ER from the

cytosol (including the ER-specific translocators themselves). The same is true for

mitochondria and plastids.

Thus, it seems that the information required to construct an organelle does not

reside exclusively in the DNA that specifies the organelle’s proteins. Information

in the form of at least one distinct protein that preexists in the organelle membrane

is also required, and this information is passed from parent cell to daughter

cells in the form of the organelle itself. Presumably, such information is essential

for the propagation of the cell’s compartmental organization, just as the information

in DNA is essential for the propagation of the cell’s nucleotide and amino acid

sequences.

As we discuss in more detail in Chapter 13, however, the ER buds off a constant

stream of transport vesicles that incorporate only a subset of ER proteins and

therefore have a composition different from the ER itself. Similarly, the plasma

membrane constantly buds off various types of specialized endocytic vesicles.

Thus, some organelles can form from other organelles and do not have to be

inherited at cell division.

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