Chapter 10: Lipids

Fatty Acids

  • Saturated Fatty Acids:
    • Lauric acid (12:0)
    • Myristic acid (14:0)
    • Palmitic acid (16:0)
    • Stearic acid (18:0)
    • Arachidic acid (20:0)
  • Unsaturated Fatty Acids:
    • Palmitoleic acid (16:1Δ9)
    • Oleic acid (18:1Δ9)
    • Linoleic acid (18:2Δ9,12)
    • Linolenic acid (18:3Δ9,12,15)
    • Arachidonic acid (20:4Δ5,8,11,14)
  • Melting Point:
    • The melting points of saturated and unsaturated fatty acids vary significantly, with saturated fatty acids generally having higher melting points. For example, saturated fatty acids can have a peak melting point of 69°C, while unsaturated fatty acids can drop as low as -16°C.

𝜔-6 and 𝜔-3 Fatty Acids

  • Naming: Fatty acids are named based on the distance of the last double bond from the methyl end of the fatty acid chain.
    • Linoleic acid: classified as 𝜔-6
    • α-linolenic acid (ALA): classified as 𝜔-3
  • Importance: Both 𝜔-6 and 𝜔-3 fatty acids are important dietary components, playing vital roles in numerous biological processes.

Triacylglycerols

  • Composition: Triacylglycerols consist of glycerol linked to three fatty acids.
  • Functions:
    • Serves as a major energy source for many organisms.
    • Most reduced form of carbon found in nature, requiring no solvation by water.
    • Allow for efficient packing of energy.
    • Storage locations:
    • Animals: mainly stored in adipose tissue
    • Plants: primarily stored in seeds
  • Additional functions include:
    • Providing insulation and water repellent properties.
    • Producing metabolic water during oxidation.
    • Formation of soap via saponification.
  • Classifications:
    • Fats: solid at room temperature; primarily composed of saturated fatty acids.
    • Oils: liquid at room temperature; primarily composed of unsaturated fatty acids.

Fatty Acids in Foods

  • General Trends:
    • Plants typically contain more unsaturated fats.
    • Animals generally contain more saturated fats.
  • Sources and Composition of fatty acids (as percentages of total fatty acids):
    • Beef: Lauric (5%), Myristic (24-32%), Palmitic (20-25%), Oleic (37-43%), Linoleic (2-3%)
    • Milk: Lauric (25%), Myristic (12%), Palmitic (33%), Oleic (3%)
    • Coconut: Lauric (74%), Myristic (10%), Palmitic (2%), Oleic (7%)
    • Palm: Lauric (39%), Myristic (4%), Palmitic (40%), Oleic (8%)
    • Corn: Lauric (8-12%), Myristic (3-4%), Palmitic (19-49%), Oleic (34-62%)
    • Olive: Lauric (9%), Myristic (2%), Palmitic (84%), Oleic (4%)
    • Safflower: Lauric (6%), Myristic (3%), Palmitic (13%), Oleic (78%)

Trans Fatty Acids

  • Formation: Trans fatty acids are formed during the hydrogenation of unsaturated fatty acids, occurring both naturally (in ruminant organisms) and synthetically.
  • Dietary Presence: They constitute a significant portion of dietary fat in the modern diet.
  • Properties:
    • Similar characteristics to saturated fatty acids.
    • Increased levels of LDL (low-density lipoproteins) and decreased levels of HDL (high-density lipoproteins).
    • Elevated triglyceride levels.
  • Examples:
    • Stearic acid
    • Oleic acid
    • Elaidic acid
  • Reasons for Partial Hydrogenation:
    • Creates a cheaper, butter-like substance.
    • Enhances cooking characteristics and stability of polyunsaturated fats.
    • Reduces oxidation sensitivity compared to monounsaturated or saturated fats.
    • Produces fatty acid aldehydes, which are generally regarded as toxic.

Waxes

  • Composition: Waxes are esters formed from long-chain alcohols and long-chain fatty acids.
  • Properties: They are highly insoluble in water.
  • Role: Waxes typically serve a protective function for leaves, fruits, skin, fur, and feathers.
  • Examples of Waxes:
    • Carnauba wax (from palm trees): consists of 80-85% wax and fatty alcohols.
    • Lanolin (from wool): contains 25% wax, along with sterol and terpene esters.
    • Beeswax: composed of 70-80% wax, ethyl palmitate, and hydrocarbons.
  • Fatty Acids in Waxes:
    • Animals: commonly palmitic or stearic acids.
    • Plants: usually unsaturated with varying chain lengths.
  • Common Alcohol: Often includes triacontanol (melissyl alcohol, 30 carbons).

Membrane Lipids

  • Types:
    • Phospholipids
    • Glycolipids
    • Glycerophospholipids
    • Sphingolipids (including Galactolipids and Sulfolipids)
  • Structural Details of Glycerophospholipids:
    • Composed of a backbone of glycerol attached to two fatty acids and a phosphate group.
    • The phosphate group is typically on the C3 hydroxyl group, while unsaturated fatty acids generally attach to the C2 position and saturated to the C1 position.

Prochirality of Glycerol

  • C-2 of Glycerol: Functions as a prochiral center, allowing distinctions into pro-(R) and pro-(S) positions:
    • Pro-(S) Position: Given higher priority (indicated by the sn prefix).
    • Example: sn-glycerol-3-phosphate indicates pro-(S) takes precedence.

Glycerophospholipids

  • Role: They are the predominant membrane lipids in bacterial cells, organelles, and the brain, especially in mitochondria.
  • Regarding Esterification: The process is extremely common and crucial for membrane formation.

Cardiolipin Variants

  • Found predominantly in bacterial and mitochondrial membranes.
  • Differences by species:
    • Yeast: Composed of solely monounsaturated fatty acids (16/18 carbon chains).
    • Mammals: Primarily consist of 18-carbon chains, with various unsaturated forms.
    • Bread Mold (N. crassa): Features mixtures of 16/18C saturated, mono-, and polyunsaturated forms.

Ether Glycerophospholipids

  • Composition Change: An ether replaces the ester at C-1 or C-2 of glycerol, a configuration more common in cardiac tissues.

Plasmalogens

  • Presence: Found in the membranes of all cells, featuring a phosphate ester with choline, ethanolamine, or serine.
  • Significance: The enol ether structure is susceptible to oxidation, which may confer antioxidant properties.

Platelet Activating Factor

  • Functions include:
    • Promotion of platelet aggregation.
    • Induction of blood vessel dilation.
    • Acting as an inflammatory mediator in egg implantation.
    • Notably water-soluble and effective at picomolar concentrations.

Unique Archaeal Membrane Lipids

  • Adaptations to extreme environments (e.g., hot springs, thermal vents) include:
    • Ether linkages instead of esters for increased stability against hydrolysis.
    • The particular structure promotes a rigid yet packed membrane structure, enhancing overall stability.

Sphingolipids

  • Membrane lipids present in all mammalian cells characterized by the presence of sphingosine, an 18-carbon amino alcohol.
  • Formation Process: Adding a fatty acid to sphingosine yields a ceramide, which can further incorporate carbohydrates to produce neutral glycolipids.

Glycosphingolipids

  • These are sphingolipids linked to carbohydrates via a 𝛽-glycosidic bond, with ceramide as the foundation.
  • ABO Blood Type Antigens:
    • Composition examples: Galactose, Glucose, Fucose, N-Acetylgalactosamine define the various antigens present.

Phospholipases

  • Enzymes critical for recycling phospholipid components, vital for membrane maintenance.
  • Example of a toxic source: Rattlesnake venom contains phospholipase A2, which disrupts extracellular leaflet membrane lipids, particularly at neuromuscular junctions, leading to membrane rupture.

Additional Lipid Classes

  • Classification: Includes
    • Isoprenoids
    • Derived from isoprene (2-methyl-1,3-butadiene).
    • Terpenes
    • Classified by the number of isoprene units, e.g., monoterpenes (2 units), sesquiterpenes (3 units), diterpenes (4 units).
  • Examples:
    • Geraniol (monoterpene)
    • Farnesol (sesquiterpene)
    • Phytol (diterpene)

Vitamins Derived from Terpenes

  • Vitamin A:
    • Crucial for vision; retinal is a derivative of β-carotene oxidation in the liver.
    • Operates in conjunction with opsin to generate rhodopsin in cone and rod cells of the retina. The isomerization of retinal is essential for the phototransduction process in vision.
  • Mixed Terpenoids:
    • These involve combinations of terpene derivatives and other functional groups, such as:
    • Vitamin E: functions as an antioxidant.
    • Vitamin K: involved in the carboxylation of prothrombin.

Antioxidant Properties of Vitamin E

  • Mechanism: Vitamin E interacts with and neutralizes oxidized membrane lipids, with Vitamin C acting to regenerate Vitamin E following its antioxidant activity.

Steroids

  • Cholesterol: Acts as a precursor for all other steroid compounds in animals; it plays significant roles in cellular structure and signaling through steroid hormones.
  • Steroid Hormones: Function in regulating salt balance, metabolic processes, and sexual function in animals.

Vitamin D

  • Role: Vital for calcium (Ca) and phosphorus (P) regulation; promotes the synthesis of Ca binding proteins, aiding intestinal Ca absorption.
  • Deficiency: In children can lead to rickets, a condition characterized by softened bones and skeletal deformities due to inadequate mineralization.

Eicosanoids

  • Classes:
    • Prostaglandins
    • Thromboxanes
    • Leukotrienes
  • Eicosanoids play complex roles in numerous physiological processes, sometimes exhibiting opposing effects (e.g., PGE2 acts as both a vasodilator and bronchial dilator).
  • Pathway Complexity: Includes multiple actions depending on the specific receptors engaged during response dynamics, such as allergic responses, uterine contractions, and promoting inflammation.

Comparison of 𝜔-3 and 𝜔-6 Fatty Acids

  • 𝜔-3 Products: Include resolvins and protectins that act as anti-inflammatory mediators.
  • 𝜔-6 Products: Known for being pro-inflammatory mediators.
    • Notably, high concentrations of 𝜔-3 may inhibit cyclooxygenase (COX) enzyme activity, limiting 𝜔-6 substrate use.

Dietary Ratios of 𝜔-3 to 𝜔-6 Fatty Acids

  • Historical Perspective: Humans evolved to consume a diet with an approximate 1:1 ratio of 𝜔-6 to 𝜔-3 fatty acids; an ideal ratio is considered to be around 2.5:1.
  • Contemporary Trends: The average American diet features an imbalance, reported at approximately 16:1 in favor of 𝜔-6 fatty acids.
  • Common Misconceptions: The myth regarding flaxseed; while high in 𝜔-3 (mostly ALA), it does not sufficiently provide EPA and DHA, which are more biologically active.
  • Conversion Rate: The conversion from ALA to EPA is less than 1% efficient.

Approximate Ratios of 𝜔-6 to 𝜔-3 in Foods**

  • Corn Oil: Approximately 50:1 ratio (65% PUFA)
  • Hemp Oil: Approximately 3:1 (predominantly ALA)
  • Canola Oil: Approximately 2:1 (mostly ALA)
  • Flax Oil: Approximately 1:2-3 (predominantly ALA)
  • Olive Oil: Ratios between 3-14:1
  • Animal Sources:
    • Corn-fed beef: ratio of 4:1
    • Grass-fed beef: ratio of 2:1
    • Wild Salmon: ratio of 1:11
    • Farmed Salmon: ratio of 1:1
    • Canned Tuna in water: ratio of 1:30
    • Canned Tuna in oil: ratio of 13:1