Solenoid model

Fig. 5. Solenoid model of the 30-nm filament of chromatin, where the disks represent nucleosomes and the dark line unbound DNA.

Although the higher order structure of chromatin is still not well understood, two models have been suggested and supported by experimental evidence (for references, see Felsenfeld, 1978 Chambon, 1978). One is the solenoid model of Finch and Klug (1976), obtained by supercoiling the chromatin thread, and the other model is the superbead model (Renz et al., 1977). Each may represent a different... 

Although the existence of the 30nm fiber is widely accepted, several models have been proposed for its structure (Felsenfeld and McGhee, 1986). Thoma and colleagues proposed a solenoid model, in which the nucleosomes are ordered in a spiral manner (Thoma et al, 1979). Woodcock and colleagues postulated a helical ribbon model, in which the nucleosomes are arranged in a zig-zag manner and the sheet of the zig-zag nucleosomes winds up helically to form a ribbon-like stmcture (Woodcock ef a/., 1984). 

With the demise of the uniform fiber model in 1974, it became necessary to devise other models to account for the early electron micrographs of chromatin fibers and the X-ray diffraction studies (see Ref. [1], Chapter 1). Two models appeared in 1976, and were the major contenders for consideration in 1978. The superbead model of Franke et al. [36] envisioned the chromatin fiber as a compaction of multi-nucleosome superbeads . The solenoid model of Finch and Klug [37] postulated a regular helical array of nucleosomes, with approximately six nucleosomes per turn and a pitch of 10 nm. Although a number of competing helical models appeared in the 1980s (see Ref. [1], Chapter 7) the solenoid model remains a serious contender to this day. Structural details of this model, such as the precise disposition of linker DNA, are still lacking. 

The structure of the condensed chromatin fiber is still under discussion [1,23,54], with two competing models the original solenoid model of Finch and Klug [16], and the straight-linker model [12,14,55]. Assessing the structure in vivo or in situ has proven impossible thus far, due to technical limitations. Chromatin fibers released from nuclei into solution by nuclease treatment have been widely used as models for fiber structure such fibers are extended at low ionic strength and condensed at ionic strengths believed to be close to those found in vivo ( 150 mM Na" " or 0.35 mM Mg " "). The salt-induced fiber compaction has been extensively studied in the past but is still poorly understood in terms not only of the details of the structure but also in terms of the molecular mechanisms of the compaction process. 

Fig. 1. The two competing classes of models for the 30-nm fiber can be distributed into (a) solenoid models and (b) crossed-linker models. For both fiber types the side and the top view are shown. Two nucleosomes that are directly connected via DNA linker are shaded in grey. In the solenoid these nucleosomes are located on the same side of the fiber requiring the linker to be bent. In the crossed-linker case they sit on opposite sides of the fiber and are connected via a straight linker.

Computational modeling can be a very powerful tool to understand the structure and dynamics of complex supramolecular assemblies in biological systems. We need to sharpen the definition of the term model somewhat, designating a procedure that allows us to quantitatively predict the physical properties of the system. In that sense, the simple geometrical illustrations in Fig. 1 only qualify if by some means experimentally accessible parameters can be calculated. As an example, a quantitative treatment of DNA bending in the solenoid model would only be possible if beyond the mechanical and charge properties of... 

A FIGURE 10-21 Solenoid model of the 30-nm condensed chromatin fiber in a side view. The octameric histone core (see Figure 10-20) is shown as an orange disk. Each nucieosome associates with one HI moiecuie, and the fiber coiis into a soienoid structure with a diameter of 30 nm. [Adapted from M. Grunstein, 1992, Sci. Am. 267 68.]... 

Currently the solenoid model and the model of Bordas et al. provide descriptions for the structure of the 30 nm fibre which are consistent with most of the observations. The parameters of the solenoid and of the model of Bordas et al. are similar. The two models differ in the path of the linker DNA in the former the linker connects the adjacent nucleosomes but in the latter nucleosomes which are not packed adjacently are connected. 

Figure 21.1 Structure of a single nucleosome (a) and proposed solenoid model of packing of the nucleosomes (b)...

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