cell membrane article notes
Lipid Bilayer as the Basis for Cell Membrane Structure
The lipid bilayer is universally established as the foundational structure of cell membranes.
Its presence is easily observable using electron microscopy.
Revealing details of its organization requires specialized techniques like:
X-ray diffraction.
Freeze-fracture electron microscopy.
The spontaneous formation of bilayers is due to the unique properties of lipid molecules under simple artificial conditions.
Membrane Lipids Are Amphipathic Molecules
Lipid molecules, primarily fatty molecules, constitute about 50% of the mass of most animal cell membranes.
Nearly all remaining mass is accounted for by proteins.
Approximately 5 × 10^6 lipid molecules can be found in a 1 μm × 1 μm area of a lipid bilayer.
This translates to nearly 10^9 lipid molecules in the plasma membrane of a small animal cell.
All lipid molecules in cell membranes are described as amphipathic (or amphiphilic):
Hydrophilic ("water-loving") or polar end.
Hydrophobic ("water-fearing") or nonpolar end.
Phospholipids as the Most Abundant Membrane Lipids
The most prevalent type of membrane lipid is phospholipids:
Composed of a polar head group and two hydrophobic hydrocarbon tails.
Tails usually consist of fatty acids, ranging in length from 14 to 24 carbon atoms.
One tail typically possesses one or more cis-double bonds (unsaturated), while the other is generally saturated.
Kinks caused by cis-double bonds affect packing and fluidity of the membrane.
Amphipathic Nature and Spontaneous Bilayer Formation
The shape and amphipathic property of lipid molecules lead to spontaneous bilayer formation in aqueous environments:
Hydrophilic molecules dissolve in water due to favorable electrostatic interactions or hydrogen bonds.
Hydrophobic molecules are insoluble; they reorganize surrounding water into less favorable structures, increasing free energy.
Burying hydrophobic tails reduces water interaction:
Aggregation can form spherical micelles or bimolecular sheets (bilayers).
Phospholipid bilayers are energetically favorable due to head group-water interaction.
Self-Healing Properties of Bilayers
Phospholipid bilayers exhibit self-healing properties:
A small tear leads to an energetically unfavorable free edge with water.
Lipids spontaneously rearrange to eliminate the free edge, forming a sealed compartment.
Larger tears are repaired via intracellular vesicle fusion.
Fluidity of Lipid Bilayers
The lipid bilayer functions as a two-dimensional fluid:
Individual lipid molecules can diffuse freely within bilayers.
Important studies conducted in the 1970s demonstrated lipid mobility within synthetic bilayers.
Experimental Setups
Two prominent preparations for studying lipid bilayers include:
Spherical vesicles (liposomes): Diameter ranging from 25 nm to 1 μm.
Planar bilayers (black membranes): Formed across holes between aqueous compartments.
Lipid Mobility Studies
Various techniques such as electron spin resonance (ESR) spectroscopy track lipid molecule motion:
Flip-flop (migration between monolayers) occurs infrequently (~< once per month).
Lateral diffusion occurs rapidly (~10^7 times per second); diffusion coefficient approximately 10^-8 cm²/sec.
Composition Impacting Fluidity
The fluidity of a lipid bilayer is significantly influenced by its composition and temperature:
Phase transition is noted when a bilayer changes from liquid to crystalline state.
The transition point is lower when hydrocarbon chains are shorter or unsaturated.
Cholesterol's Role in Membrane Fluidity
Many eukaryotic membranes consist of phospholipids and cholesterol:
Cholesterol molecules can reach ratios of 1:1 with phospholipids and enhance permeability barrier properties.
They position themselves near polar head groups and interact with regions of hydrocarbon chains to reduce mobility and permeability to small water-soluble molecules.
Cholesterol's presence inhibits crystallization of hydrocarbon chains, affecting phase transitions.
Lipid Composition Variations
Biological membranes vary in lipid composition:
Eukaryotic membranes: Large amounts of cholesterol plus various different phospholipids.
Bacterial membranes: Usually no cholesterol; stability is provided by an exterior cell wall.
Major Phospholipids in Mammalian Cells
Key phospholipids in many mammalian plasma membranes include:
Phosphatidylcholine
Phosphatidylethanolamine
Phosphatidylserine (carries a negative charge)
Sphingomyelin (electrically neutral at physiological pH).
Together, these phospholipids account for over 50% of the total lipid mass in membranes.
Functional Asymmetry of the Lipid Bilayer
The lipid bilayer exhibits asymmetry, which is functionally significant:
Composition differs between inner and outer monolayers of membranes.
For instance, in human red blood cells:
Choline-containing lipids predominantly occupy the outer monolayer.
Phosphati-dylethanolamine and phosphatidylserine exist mostly in the inner monolayer, leading to a charge difference.
Role of Lipid Asymmetry in Membrane Function
Asymmetry is crucial for cytosolic proteins that bind to specific lipid head groups.
For example, protein kinase C (PKC) binds to the cytosolic face of the plasma membrane and is activated by phosphatidylserine.
Phosphatidylinositol is also significant in cell signaling:
It can be phosphorylated to create recruiting sites for signaling proteins.
Glycolipids
Glycolipids are found exclusively in the noncytosolic monolayer of plasma membranes:
Generally represent about 5% of lipid molecules in the outer monolayer, primarily in lipid rafts.
They exhibit hydrogen bond formation between their sugar groups and van der Waals forces among hydrocarbon chains.
Types include gangliosides, which possess one or more negatively charged sialic acid residues, crucial in nerve cell membranes.
Functions of Glycolipids
Provide protection against harsh conditions in epithelial cell membranes.
Alter electrical fields and ion concentrations across membranes.
Involved in cell-recognition processes via carbohydrate-binding proteins (lectins).
Certain glycolipids, such as gangliosides, serve as entry points for toxins (e.g., cholera toxin).
Summary
Biological membranes consist of a double layer of lipid molecules, with membrane proteins embedded within.
Lipid bilayers are fluid, allowing rapid diffusion of lipid molecules within their monolayers.
Major classes of membrane lipids include phospholipids, cholesterol, and glycolipids, each contributing uniquely to membrane functionality and integrity.
The distinct lipid compositions between the inner and outer monolayers reflect their specific roles and interactions within cellular processes.