Halestrap A P
Biochem J. 1976 May 15;156(2):193-207. doi: 10.1042/bj1560193.
The kinetics and activation energy of entry of pyruvate and lactate into the erythrocyte were studied at concentrations below 4 and 15mM respectively. The Km and Vmax. values for both substrates are reported, and it is shown that pyruvate inhibits competitively with respect to lactate and vice versa. In both cases the Km for the carboxylate as a substrate was the same as its Ki as an inhibitor. Alpha-Cyano-4-hydroxycinnamate and its analogues inhibited the uptake of both lactate and pyruvate competitively. Inhibition was also produced by treatment of cells with fluorodinitrobenzene but not with the thiol reagents or Pronase. At high concentrations of pyruvate or lactate (20mM), uptake of the carboxylate was accompanied by an efflux of Cl-ions. This efflux of Cl- was inhibited by alpha-cyano-4-hydroxycinnamate and picrate and could be totally abolished by very low (less than 10 muM) concentrations of the inhibitor of Cl- transport, 4,4'-di-isothiocyanostilbene-2,2'-disulphonic acid. This inhibitor titrated out the chlordie efflux induced by pyruvate, bicarbonate, formate and fluoride, in each case total inhibition becoming apparent when approximately 1.2x10(6) molecules of inhibitor were present per erythrocyte, that is, about one inhibitor molecule per molecule of the Cl- carrier. Evan when Cl- efflux was totally blocked pyruvate and lactate uptake occurred. Kinetic evidence is presented which suggests that the Cl- carrier can transport pyruvate and lactate with a high Km and high Vmax., but that an additional carrier with a low Km and a low Vmax. also exists. This carrier catalyses the exchange of small carboxylate anions with intracellular lactate, is competitively inhibited by alpha-cyano-4-hydroxycinnamate and non-competitively inhibited by picrate. The Cl- carrier shows a reverse pattern of inhibition. It is concluded that net efflux of lactic acid from the cell must occur on the Cl- carrier and involve exchange with HCO3 - followed by loss of CO2. The low Km carrier might be used in pyruvate/lactate or acetoacetate/beta-hydroxybutyrate exchanges involved in transferring reducing power across the cell membrane. The possibility that the Cl- carrier exists in cells other than the erythrocyte is discussed. It is concluded that its presence in other cell membranes together with a low intracellular Cl- concentration would explain why the pH in the cytoplasm is lower than that of the blood, and why permeable carboxylate anions do not accumulate within the cell when added from outside.
分别在丙酮酸和乳酸浓度低于4mM和15mM的情况下,研究了它们进入红细胞的动力学和活化能。报告了两种底物的米氏常数(Km)和最大反应速度(Vmax)值,结果表明丙酮酸对乳酸的摄取具有竞争性抑制作用,反之亦然。在这两种情况下,羧酸盐作为底物的Km与其作为抑制剂的抑制常数(Ki)相同。α-氰基-4-羟基肉桂酸及其类似物竞争性抑制乳酸和丙酮酸的摄取。用氟二硝基苯处理细胞也会产生抑制作用,但用硫醇试剂或链霉蛋白酶处理则不会。在高浓度的丙酮酸或乳酸(20mM)下,羧酸盐的摄取伴随着氯离子的外流。α-氰基-4-羟基肉桂酸和苦味酸盐可抑制这种氯离子外流,极低浓度(小于10μM)的氯离子转运抑制剂4,4'-二异硫氰基芪-2,2'-二磺酸可完全消除这种外流。这种抑制剂可滴定出由丙酮酸、碳酸氢盐、甲酸盐和氟化物诱导的氯离子外流,在每种情况下,当每个红细胞中约有1.2×10⁶个抑制剂分子时,即每分子氯离子载体约有一个抑制剂分子时,完全抑制作用变得明显。即使氯离子外流被完全阻断,丙酮酸和乳酸的摄取仍会发生。动力学证据表明,氯离子载体可以高Km和高Vmax转运丙酮酸和乳酸,但也存在另一种低Km和低Vmax的载体。这种载体催化小羧酸盐阴离子与细胞内乳酸的交换,被α-氰基-4-羟基肉桂酸竞争性抑制,被苦味酸盐非竞争性抑制。氯离子载体表现出相反的抑制模式。得出的结论是,细胞内乳酸的净外流必定发生在氯离子载体上,且涉及与HCO₃⁻的交换,随后是CO₂的丢失。低Km载体可能用于丙酮酸/乳酸或乙酰乙酸/β-羟基丁酸的交换,参与跨细胞膜传递还原力。文中讨论了氯离子载体是否存在于红细胞以外的其他细胞中的可能性。得出的结论是,它在其他细胞膜中的存在以及细胞内低氯离子浓度可以解释为什么细胞质中的pH低于血液中的pH,以及当从外部添加可渗透的羧酸盐阴离子时它们为何不会在细胞内积累。