Evidence that glutamine, not sugar, is the major energy source for cultured HeLa cells.

The chromatographic identification of some biologically important phosphate esters.

Transphosphatidylation by Phospholipase D

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I i i i i i i i i

Improved Techniques in Paper Chromatography of Carbohydrates

Investigation of the reduction of carbon dioxide in plants requires a rapid and general method for separating and identifying a large number of compounds. Paper chromatography is such a method. This paper constitutes a report of progress in the development of experimental procedures in the application of paper chromatography to the separation of compounds formed by plants during short periods of photosynthesis. The compounds involved are important intermediates of plant and animal metabolism. These methods should find application not only in other biochemical problems, but in the control of chemical synthesis2 as well. A number of products formed during a short period of photosynthesis with radioactive carbon dioxide have been r e p ~ r t e d . ~ ~ * ~ ~ , ~ ~ ~ The classical biochemical methods and ion exchange techniques which were used in those identifications would require considerable effort to separate the large number of compounds involved. The success of paper chromatography in separating two groups of compounds important in phytosynthesis, amino acidss and s u g ~ t r s , ~ ~ ~ ~ prompted the present work on separation of polycarboxylic acids and phosphate esters. These compounds constitute the presently known intermediates in synthesis of plant material. l1 The primary requisite for the use of this method is some means of determining the position of a compound in the chromatogram. Both chemical and physical methods have been used. The former includes color tests which depend on development of a color directly on the paper by the use of specific reagents: ninhydrin for amino acids,*naphthoresorcinolor Tollens reagent for sugars,’” ferric ion for hydroxamic acid^,'^^^^ $H in-

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Benson, A. A., Et Al.

Papers

The path of carbon in photosynthesis. V. Paper chromatography and radioautography of the products.