¥ Figue 16, Chromatin Packing in a Eukaryotic Chromosome This setes of dagrams and transmission electron micrographs depicts a crtent model for the progress levels of DNA cong and folding. The lustranon zooms out fom single molecule of DNA toa metaphase chromosome, which sarge encugh tobe seen th a igh microscope Sn nen ¢ Se 4 i NuiclearSme ‘10 rnm ip diameter xa ‘double hex 2am in clameten Histo‘ tail Fistnes Nucleosomes, or “beads on DNA, the double helix Histones a string” (10-nm fibre) Shown here isa iblon model of DNA, Proteins called histonesare responsite for In elecvon micrographs, unfolded chromatin is ‘with each idbon representing one ofthe fist level of DNA packing in chromatin 10 ne in diameter (the 10-nm fibre). Such ‘the sugar-phosphate backbones. As Although each histone is small—conaining __—_chrornatinresemiles beads on a sting (see the you wll ecall rom Figure 16.7, the only about 100 amino acids—the total mass TEM). Each “bead isa nucleosome, the basic hosphate groups long the backbone of histone in chromatin approximately equals unit of DNA packing; the “string” between beads ‘contribute a negative charge along the the mass of DNA. More than a fith of a is called fnker DNA. ‘outside of each strand. The TEM shows _histone’s amino acids are positively charged ‘A nucleosome consisis of DNA wound twice ‘molecule of naked DNA; the double sine or arginine) and therafore bind tightly around a protein core composed of two: helix alone is 2 nm across ‘o the negatively charged DNA, imlecules of each of the four main histone types. Four types of histones are most common The amino end (N-terminus) of each histone in chromatin: H2A, 128, 13, and H4. The (the histone tai) extends outward from the histones ace very similar among eukaryotes, nucleosome. ‘or example, all but two of the amino acidsin In the cell cycle, the histones leave the DNA, ‘cow HA are identical o those in pea H4. The only briefly during DNA replication. Generally, -apoarent conservation of histone genes during they do the same during transcription, another ‘evolution probadly reflects the important role process that requires access to the DNA by the fofhtonesin oroenizng DNA within cel, cal’s molecular machinery Chapter 18 val The fourrman types of histones are citcal _cscutssorme recent nsings about the role of to thenex{lovel ofONA packing (Aft ype histone falls and nucleosomesin the regulation ofhistone called HI isimelvedin sfusher of gene expression 346 UNIT THREE Geneaes cr ee Each eukaryotic chromosome contains a singe hear Inthe cell, eukaryotic DNA is precisely combined with DNA donile helix hain humans averages about 1.5% 10° | rge amount of protein. Together, ths complex of DNA and snncleotide pairs. This isan enormous amount of DNA yrotsn, called chromatin, ts nm the muceus through an relative to chromosome’s condensed length [completely | daborat multilevel system of packing. Our current view ofthe strerched out, sich a DNA molecule wot he abot Fem sieeve vale of DNA packing in a hromneomte eine long, thousands of times the diametcr of cll mules — in Fire 16.29, Soy tis gone carcily boos reading firth and that’s not even considering the DNA of the other 45 (Chromatin undergoes striking changes in its degree of human chromosomes! yacking during the couse of the clleycle see Figure 12.7) Chyomatia oo ne) Pies en. Lem sealfad 30-nm fibre The next level of packing results from interactions benveen the histone tals of ‘one nucleosome and the linker DNA and — Logped domains Co. to 11200 wm) rucleosomes on ether side. ith i Histone, Hi, fied at this eval hase (300-nm fibre)___ Metaphase chromosome interactions cause the extended 10-nm fibre The 30am fibre, in turn, formsloaps na mitotic chremnesome, the laaped damains to call or fold, forming a chromatin fibre caled oped domains attached to a themsdves clan fldin @ manner not yet fly ‘oughly 30 nmin thickness the 301m chromosome scafold made of proteins, _understcod, further compacting al the chromatin fibre G00 TEM). Although the 30-rm fibre mgking upa 300-nm fibre Gee TEM). The to proce the characteristic metachasechromc- is quite prevalent in the interphase scatfold is rch in one ype of topetsomar-- some shown inthe micrograph above. The width ruckus, the packing atangement of ‘ase, and #1 molecules abo appear to be of one chroatidis 700 rm. Paticuar genes ruceosomes inthis orm of chromatin is present. ‘aways end up lected atthe same places in stil a matter of some debate rmetaphae chromosomes, inccating thatthe packing steps ae hihly specific and precise, CHAPTER 16 The Molecular Dass of ncritince 347