Carlson R D
Basic Life Sci. 1984;27:47-72. doi: 10.1007/978-1-4899-0375-4_3.
Neutron scattering experiments have shown that both the (H3-H4)2 and H2a-H2b histone complexes are quite asymmetric in solution. The (H3-H4)2 tetramer is an oblate or flattened structure, with a radius of gyration almost as large as that of the core octamer. If the tetramer is primarily globular, it must have an axial ratio of about 1:5. It is more likely, however, that this asymmetry results in part from N-terminal arms that extend outward approximately within the major plane of the particle. If this is the case, less asymmetric models for the globular part of the tetramer, including a dislocated disk of the type proposed by Klug et al. (23), can be made consistent with the scattering data. The H2a-H2b dimer, on the other hand, is an elongated structure. The low resolution data are in good agreement with those calculated for a cylindrical model 64 X 27 A, but other elongated models fit those data almost as well, including one that approximates free N-terminal arms at each end. Free arms are not necessary, but they must extend from the ends if they exist. A contrast matching experiment done with 50% deuterated H2b and undeuterated H2a in the reconstituted dimer showed that these two histones must each be rather elongated within the complex and are not just confined to one end. The amount of scattering contrast between the undeuterated and 50% deuterated histones was sufficient to suggest further experiments using complexes reconstituted from mixtures of undeuterated and partially deuterated histones which will help elucidate their arrangement within the histone complexes and within the octamer core of the nucleosome core particle.
中子散射实验表明,(H3-H4)2和H2a-H2b组蛋白复合物在溶液中都相当不对称。(H3-H4)2四聚体是扁球形或扁平结构,其回转半径几乎与核心八聚体的回转半径一样大。如果四聚体主要是球形的,那么它的轴比必须约为1:5。然而,这种不对称性更有可能部分是由向外延伸至颗粒主平面内的N端臂导致的。如果是这种情况,那么对于四聚体球状部分的不对称性较小的模型,包括Klug等人(23)提出的那种错位盘状模型,就可以与散射数据相一致。另一方面,H2a-H2b二聚体是一种细长结构。低分辨率数据与为64×27 Å的圆柱形模型计算的数据吻合良好,但其他细长模型对这些数据的拟合效果也几乎一样好,包括一种在两端近似自由N端臂的模型。自由臂并非必需,但如果存在,它们必须从两端延伸出来。在重组二聚体中用50%氘代的H2b和未氘代的H2a进行的对比匹配实验表明,这两种组蛋白在复合物中各自必须相当细长,而不仅仅局限于一端。未氘代和50%氘代组蛋白之间的散射对比度足以表明,可以使用由未氘代和部分氘代组蛋白混合物重组的复合物进行进一步实验,这将有助于阐明它们在组蛋白复合物以及核小体核心颗粒的八聚体核心内的排列方式。