Felix J S, DeMars R
Proc Natl Acad Sci U S A. 1969 Feb;62(2):536-43. doi: 10.1073/pnas.62.2.536.
Humans with the Lesch-Nyhan syndrome have an X-chromosomal mutant gene that causes severe neurological and developmental abnormalities. The patients are deficient in hypoxanthine-guanine phosphoribosyltransferase, which converts hypoxanthine to inosinic acid, a major precursor of adenine and guanine nucleotides. Paradoxically, the enzyme defect causes hypernormal de novo synthesis of inosinic acid, which manifests itself as excesses of hypoxanthine, xanthine, and uric acid. The first step in the de novo pathway is thought to be rate-limiting, due to feedback repression by adenine and guanine nucleotides. The derepressed rate of purine production in mutants and their failure to thrive could result from reduction in the amounts of nucleotides derived from inosinic acid to levels that are inadequate for normal feedback control and for nucleic acid synthesis needed in growth. Studies with cultured cells, reported here, support the interpretation that mutants are, in effect, nucleotide-deficient. Skin fibroblasts from patients fail to proliferate in media that do not contain supplementary adenine or folic acid, a participant in two stages of purine biosynthesis. The folic acid requirement of mutant cells is at least 50-fold greater than that of normal cells, which can synthesize all the nucleotides needed for growth without exogenous adenine. Both folic acid and adenine supplements are thought to provide mutant cells with the means of making more inosinic acid available for conversion to adenine and guanine nucleotides. It is not clear why the availability of inosinate or its conversion to other nucleotides is impaired. Therapy with adenine or folic acid begun at the time of birth may avert development of the disease in mutant males.The relevant gene is X-linked and shows clonal, single-allele-expression: phenotypically normal and phenotypically mutant clones have been derived from females heterozygous for the mutant gene. The phenotypically mutant heterozygous clones have the same requirement for adenine or folic acid as cells from hemizygous mutant males, an indication that the normal allele is repressed in these clones. The adenine-folic acid requirement of mutant cells provides a method of direct, clonal selection for rare, phenotypically normal cells in mutant populations, which is applicable to the single-active-X problem and other in vitro genetic studies.
患有莱施-奈恩综合征的人类具有一个X染色体突变基因,该基因会导致严重的神经和发育异常。患者缺乏次黄嘌呤-鸟嘌呤磷酸核糖转移酶,该酶可将次黄嘌呤转化为肌苷酸,而肌苷酸是腺嘌呤和鸟嘌呤核苷酸的主要前体。矛盾的是,这种酶缺陷会导致肌苷酸的从头合成异常活跃,表现为次黄嘌呤、黄嘌呤和尿酸过量。由于腺嘌呤和鸟嘌呤核苷酸的反馈抑制作用,从头合成途径的第一步被认为是限速步骤。突变体中嘌呤产生的去抑制速率以及它们发育不良的情况,可能是由于源自肌苷酸的核苷酸量减少到不足以进行正常反馈控制和生长所需核酸合成的水平所致。本文报道的对培养细胞的研究支持了这样一种解释,即突变体实际上是核苷酸缺乏的。患者的皮肤成纤维细胞在不含补充腺嘌呤或叶酸(嘌呤生物合成两个阶段的参与者)的培养基中无法增殖。突变细胞对叶酸的需求至少比正常细胞高50倍,正常细胞无需外源性腺嘌呤就能合成生长所需的所有核苷酸。叶酸和腺嘌呤补充剂都被认为能为突变细胞提供更多可转化为腺嘌呤和鸟嘌呤核苷酸的肌苷酸。目前尚不清楚为何肌苷酸的可用性或其向其他核苷酸的转化会受损。在出生时开始用腺嘌呤或叶酸进行治疗,可能会避免突变男性患上该疾病。相关基因是X连锁的,表现为克隆性单等位基因表达:从突变基因杂合的女性中已获得表型正常和表型突变的克隆。表型突变的杂合克隆对腺嘌呤或叶酸的需求与半合子突变男性的细胞相同,这表明正常等位基因在这些克隆中受到抑制。突变细胞对腺嘌呤-叶酸的需求为在突变群体中直接克隆选择罕见的表型正常细胞提供了一种方法,这适用于单活性X问题及其他体外遗传学研究。