Cell and Molecula…

018.04.BIO201.4B9… MORENO, DANI…
8.5: Membran...

8.5

Membrane Biosynthesis in
the Endoplasmic Reticulum
LEARNING OBJECTIVE

Describe the role of the endoplasmic reticulum in membrane biosynthesis.

Membranes do not arise de novo, that is, as new entities from pools of protein and lipid that mix
together. Instead, membranes arise from preexisting membranes. Membranes grow as newly
synthesized proteins, and lipids are inserted into existing membranes in the endoplasmic reticulum
(ER). As will be apparent in the following discussion, membrane components move from the ER to
virtually every other compartment in the cell. As the membrane moves from one compartment to the
next, its proteins and lipids are modified by enzymes that reside in the cell’s various organelles. These
modifications contribute to giving each membrane compartment a unique composition and distinct
identity.
Keep in mind that cellular membranes are asymmetric: The two phospholipid layers (leaflets) of a
membrane have different compositions. This asymmetry is established initially in the endoplasmic
reticulum. Asymmetry is maintained as membrane carriers bud from one compartment and fuse to the
next. As a result, domains situated at the cytosolic surface of the ER membrane can be identified on the
cytosolic surface of transport vesicles, the cytosolic surface of Golgi cisternae, and the internal
(cytoplasmic) surface of the plasma membrane (Figure 8.14). Similarly, domains situated at the
luminal surface of the ER membrane maintain their orientation and are found at the external
(exoplasmic) surface of the plasma membrane. In fact, in many ways, including its high calcium
concentration and abundance of proteins with disulfide bonds and carbohydrate chains, the lumen of
the ER (as well as other compartments of the secretory pathway) is a lot like the extracellular space.

FIGURE 8.14

Maintenance of membrane asymmetry.

As each protein is synthesized in the rough ER, it becomes inserted into the lipid bilayer in a predictable
orientation determined by its amino acid sequence. This orientation is maintained throughout its travels in the
Concept Check 8.4: Functions of t… Concept Check 8.5: Me
endomembrane system, as illustrated in this figure. The carbohydrate chains, which are first added in the ER,
endomembrane system, as illustrated in this figure. The carbohydrate chains, which are first added in the ER,
provide a convenient way to assess membrane sidedness because they are always present Cell onand
the cisternal
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side of the cytoplasmic membranes, which becomes the exoplasmic side of the plasma membrane following
the fusion of vesicles with the plasma membrane.
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A N I M AT I O N

Cell View Animation: Endomembrane System

Maintenance of Membrane Asymmetry

Endomembrane System
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Most membrane lipids are synthesized entirely within the endoplasmic reticulum. The major
exceptions are (1) sphingomyelin and glycolipids, whose synthesis begins in the ER and is completed in
the Golgi complex; and (2) some of the unique lipids of the mitochondrial and chloroplast membranes,
which are synthesized by enzymes that reside in those membranes. The enzymes involved in the
synthesis of phospholipids are themselves integral proteins of the ER membrane with their active sites
facing the cytosol. Newly synthesized phospholipids are inserted into the half of the bilayer facing the
cytosol. Some of these lipid molecules are later flipped into the opposite leaflet through the action of
enzymes called flippases. Lipids are carried from the ER to the Golgi complex and plasma membrane as
part of the bilayer that makes up the walls of transport vesicles.
The membranes of different organelles have markedly different lipid composition (Figure 8.15a),
which indicates that changes take place as membrane flows through the cell. Several factors may
contribute to such changes (Figure 8.15b):
1. Most membranous organelles contain enzymes that modify lipids already present within a
membrane, converting one type of phospholipid (e.g., phosphatidylserine) to another (e.g.,
phosphatidylcholine) (step 1, Figure 8.15b).
2. When vesicles bud from a compartment (as in Figure 8.2a), some types of phospholipids may be
preferentially included within the membrane of the forming vesicle, while other types may be le
behind (step 2, Figure 8.15b).
3. Cells contain lipid-transfer proteins that can bind and transport lipids through the aqueous
cytosol from one membrane compartment to another (step 3, Figure 8.15b). These proteins
facilitate the movement of specific lipids from the ER to other organelles without the involvement of
transport vesicles. Lipid transfer is thought to occur at sites where the ER comes into very close
proximity to the outer membrane of other organelles.
FIGURE 8.15
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Modifying the lipid composition of membranes.
(a) Histogram indicating percentage of each of 018.04.BIO201.4B9…
three phospholipids (phosphatidylcholine, phosphatidylserine,
and sphingomyelin) in three different cellular membranes (ER, Golgi complex, and plasma membrane). The
percentage of each lipid changes gradually as membrane flows from the ER to the Golgi to the plasma
membrane. (b) Schematic diagram showing three distinct mechanisms that might explain how the
phospholipid composition of one membrane in the endomembrane system can be different from another
membrane in the system, even though the membranous compartments are spatially and temporally
continuous. (1) The head groups of phospholipids of the bilayer are modified enzymatically; (2) the membrane
of a forming vesicle contains a different phospholipid composition from the membrane it buds from; (3) lipids
can be removed from one membrane and inserted into another membrane by lipid-transfer proteins.

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