(n-6) series (which have thefirst double bond located atthe sixth carbon atom).Both linoleic acid and
α
-linolenic acid, which arethe precursors of the (n-6)and (n-3) series, respective-ly, are essential fatty acidsbecause mammals cannotsynthesize them from otherseries of fatty acids. Mam-mals lack the enzymes tointroduce double bonds atcarbon atoms before theninth carbon atom in thefatty acid chain (countingfrom the methyl end).Therefore, these essentialfatty acids must be suppliedin the diet. Subsequent de-saturation and elongationproceed only toward thecarboxyl terminus of the fatty acid.In general, fatty acids from both series can be elon-gated and desaturated. Cats, however, have reduced
∆
6desaturase and therefore cannot adequately convertlinoleic acid to arachidonic acid. Thus, arachidonicacid and linoleic acid must be supplied by the cat
’
sdiet.
3
The elongation (increase in the number of carbonatoms) and desaturation (increase in the number of double bonds) of linoleic acid and
α
-linolenic acid arecatalyzed by the same enzymes (Figure 2). However, in-terconversion between (n-3) fatty acids and (n-6) fatty acids is impossible because elongation and desaturationoccur only toward the carboxyl terminus of the fatty acid. Arachidonic acid (which is derived from the [n-6]fatty acid linoleic acid) and eicosapentaenoic acid(which is derived from the [n-3] fatty acid
α
-linolenicacid) are both fundamental components of cytoplasmicmembranes. Further metabolism of these fatty acidsleads to the generation of eicosanoids (Figure 3).Prostaglandins, thromboxanes, and leukotrienes are allderived from the metabolism of (n-3) and (n-6) fatty acids through reactions involving cyclooxygenase andlipoxygenase enzymes.Eicosanoids are important mediators of cellular reac-tions. The eicosanoids derived from arachidonic acidand eicosapentaenoic acid have different biologic ef-fects. For example, the eicosanoids that are derivedfrom eicosapentaenoic acid are in general much less po-tent inducers of inflammation than are the eicosanoidsderived from arachidonicacid. Although no conversionbetween (n-3) and (n-6) se-ries of fatty acids takesplace, inhibition and com-petition between fatty acidsof different series have beendemonstrated.
4
–
6
For exam-ple, the metabolism of
α
-linolenic acid is inhibitedby members of the linoleic-acid family: arachidonicacid,
γ
-linolenic acid, andlinoleic acid. The competi-tive equilibrium betweenlinoleate and linolenate canbe displaced in either direc-tion, and the fatty acid fa-vored in the competitiondepends on the relative lev-els of those fatty acids inthe diet.
6
An excess of (n-6) fatty acids reduces themetabolism of
α
-linolenic acid, thus possibly leading toa deficit of its metabolites, including eicosapentaenoicacid. However, the (n-3) fatty acids are much more ef-fective in inhibiting (n-6) fatty acid metabolism thanvice versa.
4
Sources
Various diets and nutritional supplements are cur-rently marketed as sources of (n-3) fatty acids for com-panion animals. The fatty acid ratios of commercialdog foods vary widely, depending on the source of fat.Diets containing safflower oil or corn oil are likely to behigh in (n-6) fatty acids and to have a ratio of (n-6) to(n-3) greater than 30:1. Also, various (n-3) fatty acidsupplements for human use are available over thecounter.
(N-3) FATTY ACID SUPPLEMENTATIONPotential Clinical Benefits
Dietary supplementation with (n-3) fatty acids may produce desirable clinical effects in dogs or cats withvarious diseases.
2,7
–
9
In addition, extrapolation fromstudies of animal models and human trials indicatesthat these fatty acids have the potential to yield variousclinical benefits (see the box). Some of the potentialbenefits for cancer patients relate to the ability of (n-3)fatty acids to decrease production of certain cytokinesthat mediate cancer cachexia.Because of their ability to modify eicosanoid produc-tion, (n-3) fatty acids have the potential to alter func-
Small Animal
The Compendium
August 1996
ARACHIDONIC ACID
s
EICOSAPENTAENOIC ACID
s
FATTY ACID RATIOS
Linoleic acid 18:2 (n-6)COOHCOOHCH
3
CH
3
α
-Linolenic acid 18:3 (n-3)
Figure 1—
Polyunsaturated fatty acids of the (n-6) series(linoleic acid 18:2 [n-6]) and of the (n-3) series (
α
-linolenicacid 18:3 [n-3]). The number of carbon atoms is listed be-fore the colon and the number of double bonds after thecolon.
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