第10章細胞の増殖

If the DNA from all 46 chromosomes in a human cell nucleus were laid out end-to-end, it would measure approximately two meters; however, its diameter would be only 2 nm! Considering that the size of a typical human cell is about 10 µm (100,000 cells lined up to equal one meter), DNA must be tightly packaged to fit in the cell’s nucleus. At the same time, it must also be readily accessible for the genes to be expressed. For this reason, the long strands of DNA are condensed into compact chromosomes during certain stages of the cell cycle. There are a number of ways that chromosomes are compacted.

In the first level of compaction, short stretches of the DNA double helix wrap around a core of eight histone proteins at regular intervals along the entire length of the chromosome (Figure 10.4). The DNA-histone complex is called chromatin. The beadlike, histone DNA complex is called a nucleosome, and DNA connecting the nucleosomes is called linker DNA. A DNA molecule in this form is about seven times shorter than the double helix without the histones, and the beads are about 10 nm in diameter, in contrast with the 2-nm diameter of a DNA double helix.

The second level of compaction occurs as the nucleosomes and the linker DNA between them coil into a 30-nm chromatin fiber. This coiling further condenses the chromosome so that it is now about 50 times shorter than the extended form.

In the third level of compaction, a variety of fibrous proteins is used to “pack the chromatin.” These fibrous proteins also ensure that each chromosome in a non-dividing cell occupies a particular area of the nucleus that does not overlap with that of any other chromosome (see the top image in Figure 10.3).

DNA replicates in the S phase of interphase, which technically is not a part of mitosis, but must always precede it. After replication, the chromosomes are composed of two linked sister chromatids. When fully compact, the pairs of identically packed chromosomes are bound to each other by cohesin proteins. The connection between the sister chromatids is closest in a region called the centromere. The conjoined sister chromatids, with a diameter of about 1 µm, are visible under a light microscope. The centromeric region is highly condensed and thus will appear as a constricted area.

Prokaryotes have a single circular chromosome composed of double-stranded DNA, whereas eukaryotes have multiple, linear chromosomes composed of chromatin wrapped around histones, all of which are surrounded by a nuclear membrane. The 46 chromosomes of human somatic cells are composed of 22 pairs of autosomes (matched pairs) and a pair of sex chromosomes, which may or may not be matched. This is the 2n or diploid state. Human gametes have 23 chromosomes, or one complete set of chromosomes; a set of chromosomes is complete with either one of the sex chromosomes, X or Y. This is the n or haploid state. Genes are segments of DNA that code for a specific functional molecule (a protein or RNA). An organism’s traits are determined by the genes inherited from each parent. Duplicated chromosomes are composed of two sister chromatids. Chromosomes are compacted using a variety of mechanisms during certain stages of the cell cycle. Several classes of protein are involved in the organization and packing of the chromosomal DNA into a highly condensed structure. The condensing complex compacts chromosomes, and the resulting condensed structure is necessary for chromosomal segregation during mitosis.