Ariola Florly S, Li Zaiguo, Cornejo Christine, Bittman Robert, Heikal Ahmed A
Department of Bioengineering, The Pennsylvania State University, University Park, Pennsylvania, USA.
Biophys J. 2009 Apr 8;96(7):2696-708. doi: 10.1016/j.bpj.2008.12.3922.
Cholesterol-rich, liquid-ordered (L(o)) domains are believed to be biologically relevant, and yet detailed knowledge about them, especially in live cells under physiological conditions, is elusive. Although these domains have been observed in model membranes, understanding cholesterol-lipid interactions at the molecular level, under controlled lipid mixing, remains a challenge. Further, although there are a number of fluorescent lipid analogs that partition into liquid-disordered (L(d)) domains, the number of such analogs with a high affinity for biologically relevant L(o) domains is limited. Here, we use a new Bodipy-labeled cholesterol (Bdp-Chol) derivative to investigate membrane fluidity, lipid order, and partitioning in various lipid phases in giant unilamellar vesicles (GUVs) as a model system. GUVs were prepared from mixtures of various molar fractions of dioleoylphosphatidylcholine, cholesterol, and egg sphingomyelin. The L(d) phase domains were also labeled with 1,1'-didodecyl-3,3,3',3'-tetramethylindocarbocyanine (DiI-C(12)) for comparison. Two-photon fluorescence lifetime and anisotropy imaging of Bdp-Chol are sensitive to lipid phase domains in GUVs. The fluorescence lifetime of Bdp-Chol in liquid-disordered, single-phase GUVs is 5.50 +/- 0.08 ns, compared with 4.1 +/- 0.4 ns in the presence of DiI-C(12). The observed reduction of fluorescence lifetime is attributed to Förster resonance energy transfer between Bdp-Chol (a donor) and DiI-C(12) (an acceptor) with an estimated efficiency of 0.25 and donor-acceptor distance of 2.6 +/- 0.2 nm. These results also indicate preferential partitioning (K(p) = 1.88) of Bdp-Chol into the L(o) phase. One-photon, time-resolved fluorescence anisotropy of Bdp-Chol decays as a triexponential in the lipid bilayer with an average rotational diffusion coefficient, lipid order parameter, and membrane fluidity that are sensitive to phase domains. The translational diffusion coefficient of Bdp-Chol, as measured using fluorescence correlation spectroscopy, is (7.4 +/- 0.3) x 10(-8) cm(2)/s and (5.0 +/- 0.2) x 10(-8) cm(2)/s in the L(d) and L(o) phases, respectively. Experimental translational/rotational diffusion coefficient ratios are compared with theoretical predictions using the hydrodynamic model (Saffman-Delbrück). The results suggest that Bdp-Chol is likely to form a complex with other lipid molecules during its macroscopic diffusion in GUV lipid bilayers at room temperature. Our integrated, multiscale results demonstrate the potential of this cholesterol analog for studying lipid-lipid interactions, lipid order, and membrane fluidity of biologically relevant L(o) domains.
富含胆固醇的液态有序(L(o))结构域被认为具有生物学相关性,然而,关于它们的详细知识却难以捉摸,尤其是在生理条件下的活细胞中。尽管这些结构域已在模型膜中被观察到,但在可控的脂质混合条件下,从分子水平理解胆固醇与脂质的相互作用仍然是一项挑战。此外,虽然有许多荧光脂质类似物可分配到液态无序(L(d))结构域中,但对生物学相关的L(o)结构域具有高亲和力的此类类似物数量有限。在此,我们使用一种新的硼二吡咯标记的胆固醇(Bdp-Chol)衍生物,以巨型单层囊泡(GUVs)作为模型系统,研究其在各种脂质相中的膜流动性、脂质有序性和分配情况。GUVs由不同摩尔分数的二油酰磷脂酰胆碱、胆固醇和卵鞘磷脂混合制备而成。L(d)相结构域也用1,1'-二癸基-3,3,3',3'-四甲基吲哚羰花青(DiI-C(12))进行标记以便比较。Bdp-Chol的双光子荧光寿命和各向异性成像对GUVs中的脂质相结构域敏感。在液态无序的单相GUVs中,Bdp-Chol的荧光寿命为5.50±0.08纳秒,而在存在DiI-C(12)的情况下为4.1±0.4纳秒。观察到的荧光寿命降低归因于Bdp-Chol(供体)和DiI-C(12)(受体)之间的Förster共振能量转移,估计效率为0.25,供体-受体距离为2.6±0.2纳米。这些结果还表明Bdp-Chol优先分配(K(p)=1.88)到L(o)相中。Bdp-Chol的单光子时间分辨荧光各向异性在脂质双层中以三指数形式衰减,其平均旋转扩散系数、脂质有序参数和膜流动性对相结构域敏感。使用荧光相关光谱法测量,Bdp-Chol在L(d)相和L(o)相中的平移扩散系数分别为(7.4±0.3)×10^(-8)平方厘米/秒和(5.0±0.2)×10^(-8)平方厘米/秒。将实验得到的平移/旋转扩散系数比值与使用流体动力学模型(Saffman-Delbrück)的理论预测进行比较。结果表明,在室温下,Bdp-Chol在GUV脂质双层中的宏观扩散过程中可能与其他脂质分子形成复合物。我们综合的多尺度结果证明了这种胆固醇类似物在研究生物学相关L(o)结构域的脂质-脂质相互作用、脂质有序性和膜流动性方面的潜力。