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Essential Cell Biology 5th edition

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Secretory Pathways

519

cis

Golgi

network

transport

vesicle

vacuole

cis

cisterna

medial

cisterna

trans

cisterna

trans

Golgi

network

(A)

Figure 15−26 The Golgi apparatus consists of a stack of flattened,

membrane-enclosed sacs. (A) A three-dimensional model of a Golgi stack

reconstructed from a sequential series of electron micrographs of the Golgi

apparatus in a secretory animal cell. To see how such models are assembled,

watch Movie 15.8. (B) Electron micrograph of a Golgi stack from a plant cell,

where the Golgi apparatus is especially distinct; the stack is oriented as in (A).

(C) A pita-bread model of the Golgi apparatus. Vesicles are shown as olives.

(B, courtesy of George Palade.)

(B)

(C)

200 nm

The proteins travel through the cisternae in sequence ECB5 in two E15.26/15.26

ways: (1) by

means of transport vesicles that bud from one cisterna and fuse with the

next; and (2) by a maturation process in which the Golgi cisternae themselves

migrate through the Golgi stack. Proteins finally exit from the trans

Golgi network in transport vesicles destined for either the cell surface or

another organelle of the endomembrane system (see Figure 15−19).

Both the cis and trans Golgi networks are thought to be important for

protein sorting: proteins entering the cis Golgi network can either move

onward through the Golgi stack or, if they contain an ER retention signal,

be returned to the ER; proteins exiting from the trans Golgi network

are sorted according to whether they are destined for lysosomes (via

endosomes) or for the cell surface. We discuss some examples of sorting

by the trans Golgi network later, and we present some of the methods for

tracking proteins through the secretory pathways of the cell in How We

Know, pp. 520–521.

Many of the oligosaccharide chains that are added to proteins in the ER

(see Figure 15–24) undergo further modifications in the Golgi apparatus.

On some proteins, for example, more complex oligosaccharide chains

are created by a highly ordered process in which sugars are added and

removed by a series of enzymes that act in a rigidly determined sequence

as the protein passes through the Golgi stack. As would be expected, the

enzymes that act early in the chain of processing events are located in

cisternae close to the cis face, while enzymes that act late are located

in cisternae near the trans face.

Secretory Proteins Are Released from the Cell by

Exocytosis

In all eukaryotic cells, a steady stream of vesicles buds from the trans

Golgi network and fuses with the plasma membrane in the process of

exocytosis. This constitutive exocytosis pathway supplies the plasma

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