Charron François M, Blanchard Maxime G, Lapointe Jean-Yves
Groupe d'Etude des Protéines Membranaires, Département de Physique, Université de Montréal, Québec H3C 3J7, Canada.
Biophys J. 2006 May 15;90(10):3546-54. doi: 10.1529/biophysj.105.076745. Epub 2006 Feb 24.
Detection of a significant transmembrane water flux immediately after cotransporter stimulation is the experimental basis for the controversial hypothesis of secondary active water transport involving a proposed stoichiometry for the human Na(+)/glucose cotransporter (SGLT1) of two Na(+), one glucose, and 264 water molecules. Volumetric measurements of Xenopus laevis oocytes coexpressing human SGLT1 and aquaporin can be used to detect osmotic gradients with high sensitivity. Adding 2 mM of the substrate alpha-methyl-glucose (alphaMG) created mild extracellular hypertonicity and generated a large cotransport current with minimal cell volume changes. After 20, 40, and 60 s of cotransport, the return to sugar-free, isotonic conditions was accompanied by measurable cell swelling averaging 0.051, 0.061, and 0.077 nl/s, respectively. These water fluxes are consistent with internal hypertonicities of 1.5, 1.7, and 2.2 mOsm for these cotransport periods. In the absence of aquaporin, the measured hypertonicites were 4.6, 5.0, and 5.3 mOsm for the same cotransport periods Cotransport-dependent water fluxes, previously assumed to be water cotransport, could be largely explained by hypertonicities of such amplitudes. Using intracellular Na(+) injection and Na(+)-selective electrode, the intracellular diffusion coefficient for Na(+) was estimated at 0.29 +/- 0.03 x 10(-5) cm(2) s(-1). Using the effect of intracellular alphaMG injection on the SGLT1-mediated outward current, the intracellular diffusion coefficient of alphaMG was estimated at 0.15 +/- 0.01 x 10(-5) cm(2) s(-1). Although these intracellular diffusion coefficients are much lower than in free aqueous solution, a diffusion model for a single solute in an oocyte would require a diffusion coefficient three times lower than estimated to explain the local osmolyte accumulation that was experimentally detected. This suggests that either the diffusion coefficients were overestimated, possibly due to the presence of convection, or the diffusion in cytosol of an oocyte is more complex than depicted by a simple model.
共转运体刺激后立即检测到显著的跨膜水通量,这是关于继发性主动水转运这一有争议假说的实验基础,该假说涉及一种关于人类钠/葡萄糖共转运体(SGLT1)的化学计量关系,即两个钠离子、一个葡萄糖分子和264个水分子。对共表达人类SGLT1和水通道蛋白的非洲爪蟾卵母细胞进行体积测量,可用于高灵敏度地检测渗透压梯度。添加2 mM的底物α-甲基葡萄糖(αMG)会造成轻度的细胞外高渗,并产生较大的共转运电流,而细胞体积变化最小。在共转运20、40和60秒后,恢复到无糖等渗条件时,可测量到细胞平均分别以0.051、0.061和0.077 nl/s的速率肿胀。这些水通量与这些共转运时间段内1.5、1.7和2.2 mOsm的细胞内高渗情况一致。在没有水通道蛋白的情况下,相同共转运时间段内测得的高渗值分别为4.6、5.0和5.3 mOsm。以前认为依赖共转运的水通量是水的共转运,而这种幅度的高渗情况在很大程度上可以解释这种现象。使用细胞内注射钠离子和钠离子选择性电极,估计钠离子在细胞内的扩散系数为0.29±0.03×10⁻⁵ cm² s⁻¹。利用细胞内注射αMG对SGLT1介导的外向电流的影响,估计αMG在细胞内的扩散系数为0.15±0.01×10⁻⁵ cm² s⁻¹。尽管这些细胞内扩散系数远低于自由水溶液中的扩散系数,但对于卵母细胞中单一溶质的扩散模型而言,要解释实验检测到的局部渗透溶质积累,所需的扩散系数要比估计值低三倍。这表明要么扩散系数被高估了,可能是由于对流的存在,要么卵母细胞胞质溶胶中的扩散比简单模型所描述的更为复杂。