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The Contribution of Adolf Windaus History of Nutrition The Discovery of Vitamin D: The Contribution of Adolf Windaus Downloaded from https://academic.oup.com/jn/article-abstract/134/6/1299/4688802 by Western University of Health Sciences Library user on 18 February 2020 George Wolf1 Department of Nutritional Sciences and Toxicology, University of California, Berkeley, CA 94720-3104 The Nobel prize for chemistry for 1928 was awarded to so treated no longer had growth-promoting activity; the rats Adolf Windaus “for his studies on the constitution of the fed the treated butterfat developed xerophthalmia and died sterols and their connection with vitamins” (1), the first within 40–50 d. person to receive an award mentioning vitamins. What was The key experiment was performed by McCollum and his the contribution Windaus made to our knowledge of vitamins co-workers (8) in 1922, when they observed that heated, that deserved the highest scientific accolade? oxidized cod-liver oil could not prevent xerophthalmia but The vitamin in question was vitamin D. It had a long could cure rickets in the rats. “This shows that oxidation history before Windaus appeared on the scene. Rickets, the destroys fat-soluble A without destroying another substance bone disease caused by vitamin D deficiency, was known in which plays an important role in bone growth” (8). They antiquity and was described in detail by F. Glisson in 1650 (2). concluded that fat-soluble factor A consisted of 2 entities, one Many causes and cures for rickets had been proposed. Al- later called “vitamin A,” the other being the newly discovered though cod-liver oil had been used medicinally for a long time, antirickets factor. Because the water-soluble factors then dis- D. Scheutte (2) in 1824 was the first to prescribe it for the covered were termed vitamin B and the known antiscurvy treatment of rickets. It was not until 1906 that Hopkins (3) factor was called vitamin C, they named the new factor postulated the existence of essential dietary factors necessary vitamin D. for the prevention of diseases such as scurvy or rickets. In the meantime, an entirely different cure for rickets The first scientific approach to the disease was made by appeared, in the role of UV light. A long-standing tradition McCollum and his co-workers. In their early research (4) in held that fresh air and sunshine were good for the prevention 1914, they isolated a fat-soluble, nonsaponifiable factor from of rickets. Hess and Unger (9), in 1921, put forward the butterfat, necessary for normal growth and prevention of the explanation of their clinical observations that “seasonal inci- eye disease xerophthalmia in young rats. They named this dence of rickets is due to seasonal variations of sunlight.” In factor “fat-soluble factor A,” later “vitamin A.” The notion of her work with children, Chick and her team (6), mentioned a fat-soluble essential dietary factor for health led Mellanby above, observed that sunlight would cure rickets just as well as (5) in 1919 to experiment with puppies in which he succeeded cod-liver oil. in producing a bone disease by feeding them a diet of low-fat The field received a new impetus when Huldschinsky (10) milk and bread. He diagnosed rickets by X-ray examination, in 1919 argued that, if sunlight at the seaside or in the bone-calcium assay, and histology of bone, and noted that the mountains can prevent or cure rickets, then artificial sunlight, gross appearance of the dogs was quite similar to that of simulating light at mountain heights (“Ho¨hensonne”) should rachitic children. Even adding yeast to the dogs’ diet (to do the same. He exposed severely rachitic children to irradi- provide the water-soluble B-vitamins) and orange juice (to ation with a quartz-mercury lamp (emitting UV light) every prevent scurvy), did not prevent the appearance of rickets other day for 2 to 20 min for 2 mo and observed great within 3–4 mo. Rickets was prevented by the addition of improvement, including fresh calcium deposition, as revealed butterfat to their diet or, most effectively, of cod-liver oil. He by X-rays. He was careful to make sure that the children had wrote: “Rickets is a deficiency disease which develops in con- not been exposed to sunlight or received any supplements to sequence of the absence of some accessory food factor or their diet during those months. factors. It therefore seems probable that the cause of rickets is The thinking at that point was that rickets can be pre- a diminished intake of an anti-rachitic factor, which is either vented or cured by a component of butterfat or cod-liver oil [McCollum’s] fat-soluble factor A, or has a similar distribution that was distinct from the fat-soluble factor A (vitamin A). It to it” (5). A landmark investigation was that of Hariette can also be cured by an entirely different process, by sunlight Chick and her co-workers (6) who, in 1922, working with or UV light irradiation simulating sunlight, perhaps as the malnourished children in a clinic in post-World War I Vi- result of generally improved health. enna, showed that rickets prevalent in the children could be This dichotomy was jolted by the most surprising observa- cured by whole milk or cod-liver oil. tions, made simultaneously in 1924 in 3 different laboratories. In 1920 Hopkins (7) found that the fat-soluble factor A in Hume and Smith (11,12) found that rats suffering from rickets butterfat could be destroyed by heating and aeration. Butterfat induced by a low-phosphate diet (13) benefited from irradia- tion by UV light, not only by irradiation of the rats them- selves, but also by irradiation of the “air” in the glass jars from 1 To whom correspondence should be addressed. which they had been removed and then put back after irradi- E-mail: [email protected]. ation. It turned out that it was the irradiated sawdust, feces, 0022-3166/04 $8.00 © 2004 American Society for Nutritional Sciences. J. Nutr. 134: 1299–1302, 2004. Manuscript received 12 January 2004. 1299 1300 WOLF and spilt food left in the jars, which the rats later ate, and not “the precursor of vitamin D is not cholesterol itself, but a the air that improved the rats’ rickets. substance which is associated with and follows ‘chemically Goldblatt and Soames (14) observed that livers of irradi- pure’ cholesterol in all its stages of purification by the usual ated rats were curative when fed to rachitic rats. Steenbock methods (saponification and recrystallization).” The investi- and Black (15) went a step further. They argued that because gators converted the “pure” cholesterol into its dibromide, liver in the living rat is activated by UV light, perhaps liver recrystallized this, and recovered cholesterol upon treatment Downloaded from https://academic.oup.com/jn/article-abstract/134/6/1299/4688802 by Western University of Health Sciences Library user on 18 February 2020 removed from the animals can also be so activated. Indeed, with sodium amalgam. When purified by this method, the both liver and muscle tissue from nonirradiated rats, after recovered cholesterol, upon irradiation, had completely lost its removal from the body and exposed to UV light “was found to antirachitic potency. The authors state (22): “According to have become activated, being both growth-promoting and information received from Prof. Windaus, a specimen of bone-calcifying,” when fed to nonirradiated rachitic rats. cholesterol [was] prepared by him at our suggestion [my The third laboratory that simultaneously reported the im- italics] by the same procedure. Thus purified, cholesterol parting of antirachitic properties to inert foods, such as wheat, prepared in this way was no longer rendered anti-rachitic by lettuce, or cottonseed oil, was that of Hess and Weinstock irradiation with ultraviolet light. It is evident, therefore, (16,17). They also showed that in linseed oil, the antirachitic that not cholesterol, but [another] substance is the imme- properties resided in the nonsaponifiable fraction and that diate precursor of vitamin D.” activation occurred by UV irradiation in the absence of oxy- From this report, it is clear that, in the close collaboration gen (18). among Rosenheim, Hess, and Windaus, it was Rosenheim’s The discovery that irradiation of food, in particular of team that performed the crucial experiment demonstrating whole milk (containing butterfat), could impart antirachitic that “pure” cholesterol contained an impurity that could be potency led to tremendous advances in public health. The converted into the antirachitic vitamin photochemically. This procedure, when adopted by producers, caused a rapid decline was the essential clue that led to the identification of the in the prevalence of rickets in children. vitamin D precursor or provitamin. It was well known at that time (1924) that certain animal Windaus and Hess (21) improved upon the chemical puri- fats, such as butterfat or cod-liver oil, were antirachitic with- fication of the provitamin by the debromination of the cho- out irradiation. What, then, was the substance in vegetable lesterol dibromide by zinc dust. Rosenheim and Webster (23) oils that could be activated by irradiation? In 1925, Hess and emphasized that the work resulted from mutual suggestions his team (19) isolated sitosterol (then called phytosterol) from emanating from the 3 teams in London, Go¨ttingen, and New cottonseed oil, an abundant fraction in the nonsaponifiable York simultaneously, “according to a friendly arrangement.” portion of the oil. What appeared to the investigators to be With the discovery that “pure” cholesterol contained a sitosterol was inactive against rickets in rats. Upon irradiation small amount of an impurity that appeared to be the provita- by UV light, it became active.
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