Regulation of Metabolism 667
acid). This is similar to the way the thyroid hormone receptor
(for triiodothyronine) must dimerize with the RXR receptor to
activate its target genes (chapter 11, section 11.2; see fig. 11.7).
This overlapping of receptors may permit “crosstalk” between
the actions of thyroid hormone, vitamin D, and vitamin A.
In view of this, it is not surprising that thyroxine, vitamin A, and
fig. 11.7) and directly produces the effects of vitamin A. Retinoic
acid is involved, for example, in regulating embryonic develop-
ment; vitamin A deficiency interferes with embryonic development,
while excessive vitamin A during pregnancy can cause birth defects.
Retinoic acid also regulates epithelial membrane structure and func-
tion, and has various effects on the immune system that include the
induction of regulatory T lymphocytes (chapter 15, section 15.3).
These actions may account for the effectiveness of retinoids in treat-
ing acne and other skin conditions.
Vitamin D is produced by the skin under the influence of ultra-
violet light, but usually it is not produced in sufficient amounts for
all of the body’s needs. That is why we must eat food containing
additional amounts of vitamin D, and why it is classified as a vita-
min even though it can be produced by the body. The vitamin D
secreted by the skin or consumed in the diet is inactive in its
original form; it must first be converted into a derivative (1,25-
dihydroxyvitamin D 3 —see fig. 19.21 ) by enzymes in the liver
and kidneys before it can be active in the body. Once the active
derivative is produced, vitamin D helps regulate calcium balance,
primarily by promoting the intestinal absorption of calcium.
The nuclear receptor protein for 1,25-dihydroxyvitamin D 3
cannot activate a gene unless it forms a dimer with the RXR recep-
tor, which binds to the active form of retinoic acid (9- cis retinoic
CLINICAL APPLICATION
Psoriasis is an inherited skin condition characterized by
inflammation and excessive proliferation of keratinocytes (cells
of the epidermis that produce the protein keratin). Immune
responses, particularly those of T cells, are central to the
development of psoriasis. Vitamin D, applied topically on the
skin and/or taken as an oral supplement, and ultraviolet B light,
which stimulates endogenous vitamin D production, are com-
mon psoriasis treatments because of the ability of vitamin D to
reduce inflammation. It may do this by modifying gene expres-
sion and by inducing the formation of regulatory T cells (chap-
ter 15, section 15.3). Hypercalcemia is a possible side effect of
too much oral vitamin D because of its stimulation of intestinal
calcium phosphate absorption.
Vitamin Sources Function Deficiency Symptom(s)
A Yellow vegetables and fruit Constituent of visual pigment;
strengthens epithelial membranes
Night blindness; dry skin
B 1 (Thiamine) Liver, unrefined cereal grains Cofactor for enzymes that catalyze
decarboxylation
Beriberi; neuritis
B 2 (Riboflavin) Liver, milk Part of flavoproteins (such as FAD) Glossitis; cheilosis
B 6 (Pyridoxine) Liver, corn, wheat, and yeast Coenzyme for decarboxylase and
transaminase enzymes
Convulsions
B 12 (Cyanocobalamin) Liver, meat, eggs, milk Coenzyme for amino acid metabolism;
needed for erythropoiesis
Pernicious anemia
Biotin Egg yolk, liver, tomatoes Needed for fatty acid synthesis Dermatitis; enteritis
C Citrus fruits, green leafy
vegetables
Needed for collagen synthesis in
connective tissues
Scurvy
D Fish, liver Needed for intestinal absorption of
calcium and phosphate
Rickets; osteomalacia
E Milk, eggs, meat, leafy
vegetables
Antioxidant Muscular dystrophy (nonhereditary)
Folate Green leafy vegetables Needed for reactions that transfer
one carbon
Sprue; anemia
K Green leafy vegetables Promotes reactions needed for function
of clotting factors
Hemorrhage; inability to form clot
Niacin Liver, meat, yeast Part of NAD and NADP Pellagra
Pantothenic acid Liver, eggs, yeast Part of coenzyme A Dermatitis; enteritis; adrenal insufficiency
Table 19.3 | The Major Vitamins