A Membrane Is a Lipid Bilayer with Proteins Embedded in It

When exposed to an aqueous environment, amphipathic molecules undergo hydrophobic interactions.

In a membrane, for example, phospholipids are organized into two layers: Their polar heads face
outward toward the aqueous environment on both sides, and their hydrophobic tails are hidden from
the water by interacting with the tails of other molecules oriented in the opposite direction. The
resulting structure is the lipid bilayer, shown in Figure 2-12. The heads of both layers face outward, and
the hydrocarbon tails extend inward, forming the continuous hydrophobic interior of the membrane.
Every known biological membrane has such a lipid bilayer as its basic structure. Each of the lipid layers is
typically 3–4 nm thick, so the bilayer has a width of 7–8 nm. It is the lipid bilayer that gives each
membrane its characteristic “railroad track” appearance when seen in cross section with the
transmission electron microscope (TEM; Figure 2-13a).

The Lipid Bilayer as the Basis of Membrane Structure. Because of their amphipathic nature,
phospholipids in an aqueous environment orient themselves in a double layer, with the hydrophobic
tails (zigzag lines) buried on the inside and the hydrophilic heads (orange) interacting with the aqueous
environment on either side of the membrane.

3 Membranes and Membrane Structure. (a) With the transmission electron microscope (TEM), each of
the cell membranes surrounding two adjacent cells appears as a pair of dark bands that result when
osmium interacts with the hydrophilic heads of the phospholipid molecules but not with their
hydrophobic tails. (b) Biological membranes consist of amphipathic proteins embedded within a lipid
bilayer. Proteins are positioned in the membrane so that their hydrophobic regions are located in the
hydrophobic interior of the phospholipid bilayer and their hydrophilic regions are exposed to the
aqueous environment on either side of the membrane.

The osmium used to prepare the tissue for electron The osmium used to prepare the tissue for electron
microscopy reacts with the hydrophilic heads of the phospholipid molecules at both surfaces of the
membrane but not with the hydrophobic tails in the interior of the membrane, giving each membrane
its three-layered appearance. The structure of biological membranes is illustrated in Figure 2-13b.
Embedded within or associated with the membrane lipid bilayer are various membrane proteins. These
proteins are almost always amphipathic, and they become oriented in the lipid bilayer accordingly.
Hydrophobic regions of the protein associate with the interior of the membrane, whereas hydrophilic
regions protrude into the aqueous environment on either or both surfaces of the membrane. Depending
on the particular membrane, the membrane proteins may play any of a variety of roles. Some are
transport proteins, responsible for moving specific substances across an otherwise impermeable
membrane. Others are enzymes that catalyze reactions associated with the specific membrane. Still
others are the receptors on the outer surface of the cell membrane, the electron transport
intermediates of the mitochondrial membrane, or the chlorophyll-binding proteins of the chloroplast.
We will encounter each of these kinds of membrane proteins in subsequent chapters, beginning with
Chapter 4 (see, for example, Figure 4-9).

Membranes Are Selectively Permeable

Because of its hydrophobic interior, a membrane is readily permeable to nonpolar molecules. However,
it is quite impermeable to most polar molecules and is highly impermeable to all ions. Because most
cellular constituents are either polar or charged, they have little or no a The Importance of Synthesis by
Polymerizationffinity for the membrane interior and are effectively prevented from entering or escaping
from the cell. Very small molecules are an exception, however. Compounds with molecular weights
below about 100 diffuse across membranes regardless of whether they are nonpolar ( and ) or polar
(ethanol and urea). Water is an especially important example of a very small molecule that, although
polar, diffuses rapidly across membranes and can readily enter or leave cells. In contrast, even the
smallest ions are effectively excluded from the hydrophobic interior of the membrane. For example, a
lipid bilayer is at least times less permeable to such small cations as or than it is to water. This striking
difference is due to both the charge on an ion and the sphere of hydration surrounding the ion. Of
course, it is essential that cells have ways of transferring not only ions such as , and , but also a wide
variety of polar molecules across membranes that are not otherwise permeable to these substances. As
already noted, membranes are equipped with transport proteins to serve this function. A transport
protein is a specialized transmembrane protein that serves either as a hydrophilic channel through an
otherwise hydrophobic membrane or as a carrier that binds a specific solute on one side of the K+ Na+
K+ Na+ 108 O2 CO2 membrane and then undergoes a conformational change to move the solute across
the membrane. Whether a channel or a carrier, each transport protein is specific for a particular
molecule or ion (or, in some cases, for a class of closely related molecules or ions). Moreover, the
activities of these proteins can be carefully regulated to meet cellular needs. As a result, biological
membranes are best described as selectively permeable. Except for very small molecules and nonpolar
molecules, molecules and ions can move across a particular membrane only if the membrane contains
an appropriate transport protein.
The Importance of Synthesis by Polymerization

For the most part, cellular structures such as ribosomes, chromosomes, membranes, flagella, and cell
walls are made up of ordered arrays of linear polymers that are called macromolecules. Important
macromolecules in cells include proteins, nucleic acids (both DNA and RNA), and polysaccharides such as
starch, glycogen, and cellulose. Lipids are sometimes regarded as macromolecules as well; however,
they differ somewhat from the other classes of macromolecules in the way they are synthesized and will
not be discussed further until Chapter 3. Macromolecules are important in both the structure and the
function of cells. To understand the biochemical basis of cell biology, therefore, really means to
understand macromolecules—how they are made, how they are assembled, and how they function.

Macromolecules Are Responsible for Most of the Form and Function in Living Systems.

The importance of macromolecules in cell biology is emphasized by the cellular hierarchy shown in
Figure 2-14. We use the term hierarchy here to denote how biological molecules and structures can be
organized into a series of levels, each level building on the preceding one. Most cellular structures are
composed of small, water-soluble organic molecules (level 1) that cells either obtain from other cells or
synthesize from simple nonbiological molecules such as carbon dioxide, ammonia, or phosphate ions
that are available from the environment. Small organic molecules polymerize to form biological
macromolecules (level 2) such as polysaccharides, proteins, or nucleic acids. These macromolecules may
function on their own, or they can then be assembled into a variety of supramolecular structures (level
3). These supramolecular structures are themselves components of organelles and other subcellular
structures (level 4) that make up the cell itself (level 5). One of the examples in Figure 2-14 is that of cell
wall biogenesis (panels on left). In plants, a major component of the primary cell wall (level 3) is the
polysaccharide cellulose (level 2). Cellulose is a repeating polymer of the simple monosaccharide glucose
(level 1), a sugar formed by the plant cell from carbon dioxide and water in the process of
photosynthesis.