The Energy from Fatty Acids

This finding has been replicated several times in clinical studies. Let me cite one example. In a careful metabolic study carried out in 1990, Mensink and Katan determined the plasma LDL/HDL ratio when 10% of the energy from oleic acid was replaced in the diet by either the corresponding trans fat or the corresponding saturated fatty acid, stearic acid. The resulting LDL/HDL ratios were 2.02 on the oleic acid diet, 2.34 on the stearic acid diet, and 2.58 on the trans fatty acid diet. This is one more example of the impact of small structural changes in molecules on their biological properties. 

In the second stage of fatty acid oxidation, the acetyl-CoA is oxidized to C02 in the citric acid cycle. A large fraction of the theoretical yield of free energy from fatty acid oxidation is recovered as ATP by oxidative phosphorylation, the final stage of the oxidative pathway. 

In brown fat, which is specialized for the production of metabolic heat, electron transfer is uncoupled from ATP synthesis and the energy of fatty acid oxidation is dissipated as heat. 

FIGURE 4 8 Oxidation of carbohydrate and fatty adds for the production of energy. In the resting state, glycogen in the liver is broken down to glucose units however, both liver (kft) and muscle (right) derive most of their energy from fatty acids released from adipose tissue into the bloodstrecim and oxidized via the Krebs cycle. When carbohydrate is oxidized in the liver, it tends to be released into the bloodstream as pyruvate and lactate. 

For a given physiologic environment, the heart selects the most efficient substrate for energy production. A fitting example is the switch from fatty acid to carbohydrate oxidation with an acute work jump or increase in workload.15 The transient increase in rates of glycogen oxidation is followed by a sustained increase in rates of glucose and lactate oxidation (Fig. 2). Because oleate oxidation remains unaffected by the work jump, the increase in 02 consumption and cardiac work are entirely accounted for by the increase in carbohydrate oxidation. 

Apart from the symptoms we have already discussed, type 1 diabetics tend to present with another characteristic readily picked up by an alert clinician. Since sugar is not being properly handled, the tissues have to obtain more of their energy from fatty acid oxidation, but the capacity of the Krebs cycle tends also to be impaired, so that in uncontrolled type 1 diabetes there is a pile-up of ketone bodies made by the Uver. This is discussed in more detail in Topic 21. Acetoacetic acid, one of the two major ketone bodies, spontaneously breaks down to acetone, which, being very volatile, is easily lost into the air the patient breathes out. The smell of acetone on the breath is unmistakeable. This should not of course be detectable if the diabetes is well managed. 

Fa.ts nd Oils. Eats and oils from rendering animal and fish offal and vegetable oilseeds provide nutritional by-products used as a source of energy, unsaturated fatty acids, and palatabiHty enhancement. Eats influence the texture in finished pet foods. The use and price of the various melting point fats is deterrnined by the type and appearance of the desired finished food appearance. 

Ascorbic acid is involved in carnitine biosynthesis. Carnitine (y-amino-P-hydroxybutyric acid, trimethylbetaine) (30) is a component of heart muscle, skeletal tissue, Uver and other tissues. It is involved in the transport of fatty acids into mitochondria, where they are oxidized to provide energy for the ceU and animal. It is synthesized in animals from lysine and methionine by two hydroxylases, both containing ferrous iron and L-ascorbic acid. Ascorbic acid donates electrons to the enzymes involved in the metabohsm of L-tyrosine, cholesterol, and histamine (128). 

Fiber components are the principal energy source for colonic bacteria with a further contribution from digestive tract mucosal polysaccharides. Rate of fermentation varies with the chemical nature of the fiber components. Short-chain fatty acids generated by bacterial action are partiaUy absorbed through the colon waU and provide a supplementary energy source to the host. Therefore, dietary fiber is partiaUy caloric. The short-chain fatty acids also promote reabsorption of sodium and water from the colon and stimulate colonic blood flow and pancreatic secretions. Butyrate has added health benefits. Butyric acid is the preferred energy source for the colonocytes and has been shown to promote normal colonic epitheUal ceU differentiation. Butyric acid may inhibit colonic polyps and tumors. The relationships of intestinal microflora to health and disease have been reviewed (10). 

Fujiwara et al. used the CMC values of sodium and calcium salts to calculate the energetic parameters of the micellization [61]. The cohesive energy change in micelle formation of the a-sulfonated fatty acid methyl esters, calculated from the dependency of the CMC on the numbers of C atoms, is equivalent to that of typical ionic surfactants (Na ester sulfonates, 1.1 kT Ca ester sulfonates, 0.93 kT Na dodecyl sulfate, 1.1 kT). The degree of dissociation for the counterions bound to the micelle can be calculated from the dependency of the CMC on the concentration of the counterions. The values of the ester sulfonates are also in the same range as for other typical ionic surfactants (Na ester sulfonates, 0.61 Ca ester sulfonates, 0.70 Na dodecyl sulfate, 0.66). 

In the organism tissues, fatty acids are continually renewed in order to provide not only for the energy requirements, but also for the synthesis of multicomponent lipids (triacylglycerides, phospholipids, etc.). In the organism cells, fatty acids are resynthetized from simpler compounds through the aid of a supramolecular multienzyme complex referred to as fatty acid synthetase. At the Lynen laboratory, this synthetase was first isolated from yeast and then from the liver of birds and mammals. Since in mammals palmitic acid in this process is a major product, this multienzyme complex is also called palmitate synthetase. 

Calculating energy costs for the synthesis of a CK, fatty acid from acetyl-CoA is not as simple as you might first think. The major complication is that acetyl-CoA is made in the mitochondria, but fatty acid synthesis occurs in the cytosol—acetyl-CoA can t cross the mitochondrial membrane. Acetyl-CoA gets out of the mitochondria disguised as citrate. The acetyl-CoA is condensed with oxaloacetate to give citrate, and the citrate leaves the mitochondria. In the cytosol, the citrate is cleaved by an ATP-dependent citrate lyase into acetyl-CoA and oxaloacetate ... 

The citrate shuttle transports acetyl CoA groups from the mitochondria to the cytoplasm for fatty acid synthesis. Acetyl CoA combines with oxaloacetate in the mitochondria to form citrate, but rather than continuing in the citric add cycle, citrate is transported into the cytoplasm. Factors that indirectly promote this process indude insuKn and high-energy status. 

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