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Along the top are the patterns of more Chromosomes Exist in Different States Throughout the Life of a Cell We have seen how genes are arranged in chromosomes, but to form a functional chromosome , a DNA molecule must be able to do more than simply carry genes: it must be able to replicate, and the replicated copies must be separated and reliably partitioned into daughter cells at each cell division.

This process occurs through an ordered series of stages, collectively known as the cell cycle. The cell cycle is briefly summarized in Figure , and discussed in detail in Chapter Only two of the stages of the cycle concern us in this chapter.

During interphase chromosomes are replicated, and during mitosis they become highly condensed and then are separated and distributed to the two daughter nuclei. The highly condensed chromosomes in a dividing cell are known as mitotic chromosomes.

This is the form in which chromosomes are most easily visualized; in fact, all the images of chromosomes shown so far in the chapter are of chromosomes in mitosis. This condensed state is important in allowing the duplicated chromosomes to be separated by the mitotic spindle during cell division, as discussed in Chapter Figure A simplified view of the eucaryotic cell cycle.

During interphase, the cell is actively expressing its genes and is therefore synthesizing proteins. Also, during interphase and before cell division, the DNA is replicated and the chromosomes are duplicated.

During the portions of the cell cycle when the cell is not dividing, the chromosomes are extended and much of their chromatin exists as long, thin tangled threads in the nucleus so that individual chromosomes cannot be easily distinguished Figure We refer to chromosomes in this extended state as interphase chromosomes. A comparison of extended interphase chromatin with the chromatin in a mitotic chromosome.

A An electron micrograph showing an enormous tangle of chromatin spilling out of a lysed interphase nucleus. B A scanning electron micrograph of a mitotic chromosome: more Each DNA Molecule That Forms a Linear Chromosome Must Contain a Centromere, Two Telomeres, and Replication Origins A chromosome operates as a distinct structural unit: for a copy to be passed on to each daughter cell at division, each chromosome must be able to replicate, and the newly replicated copies must subsequently be separated and partitioned correctly into the two daughter cells.

These basic functions are controlled by three types of specialized nucleotide sequence in the DNA , each of which binds specific proteins that guide the machinery that replicates and segregates chromosomes Figure Figure The three DNA sequences required to produce a eucaryotic chromosome that can be replicated and then segregated at mitosis.

Each chromosome has multiple origins of replication, one centromere, and two telomeres. Shown here is the sequence of events a typical more Experiments in yeasts, whose chromosomes are relatively small and easy to manipulate, have identified the minimal DNA sequence elements responsible for each of these functions. One type of nucleotide sequence acts as a DNA replication origin , the location at which duplication of the DNA begins.

Eucaryotic chromosomes contain many origins of replication to ensure that the entire chromosome can be replicated rapidly, as discussed in detail in Chapter 5.

After replication, the two daughter chromosomes remain attached to one another and, as the cell cycle proceeds, are condensed further to produce mitotic chromosomes. The presence of a second specialized DNA sequence, called a centromere , allows one copy of each duplicated and condensed chromosome to be pulled into each daughter cell when a cell divides.

A protein complex called a kinetochore forms at the centromere and attaches the duplicated chromosomes to the mitotic spindle , allowing them to be pulled apart discussed in Chapter The third specialized DNA sequence forms telomeres , the ends of a chromosome.

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Telomeres contain repeated nucleotide sequences that enable the ends of chromosomes to be efficiently replicated. Telomeres also perform another function: the repeated telomere DNA sequences, together with the regions adjoining them, form structures that protect the end of the chromosome from being recognized by the cell as a broken DNA molecule in need of repair. We discuss this type of repair and the other features of telomeres in Chapter 5.

In yeast cells, the three types of sequences required to propagate a chromosome are relatively short typically less than base pairs each and therefore use only a tiny fraction of the information-carrying capacity of a chromosome. Although telomere sequences are fairly simple and short in all eucaryotes, the DNA sequences that specify centromeres and replication origins in more complex organisms are much longer than their yeast counterparts.

For example, experiments suggest that human centromeres may contain up to , nucleotide pairs. It has been proposed that human centromeres may not even require a stretch of DNA with a defined nucleotide sequence; instead, they may simply create a large, regularly repeating protein - nucleic acid structure. We return to this issue at the end of the chapter when we discuss in more general terms the proteins that, along with DNA, make up chromosomes.

Recall from earlier in this chapter that human chromosome 22 contains about 48 million nucleotide pairs. Stretched out end to end, its DNA would extend about 1. This remarkable feat of compression is performed by proteins that successively coil and fold the DNA into higher and higher levels of organization.

Although less condensed than mitotic chromosomes, the DNA of interphase chromosomes is still tightly packed, with an overall compaction ratio of approximately fold.


In the next sections we discuss the specialized proteins that make the compression possible. In reading these sections it is important to keep in mind that chromosome structure is dynamic. Not only do chromosomes globally condense in accord with the cell cycle, but different regions of the interphase chromosomes condense and decondense as the cells gain access to specific DNA sequences for gene expression , DNA repair , and replication. The packaging of chromosomes must therefore be accomplished in a way that allows rapid localized, on-demand access to the DNA.

Nucleosomes Are the Basic Unit of Eucaryotic Chromosome Structure The proteins that bind to the DNA to form eucaryotic chromosomes are traditionally divided into two general classes: the histones and the nonhistone chromosomal proteins.

The complex of both classes of protein with the nuclear DNA of eucaryotic cells is known as chromatin. Histones are present in such enormous quantities in the cell about 60 million molecules of each type per human cell that their total mass in chromatin is about equal to that of the DNA.

Histones are responsible for the first and most basic level of chromosome organization, the nucleosome , which was discovered in Black can be stark, yes, but it can also be professional, clean, and even sophisticated.

Pink — besides the obvious feminine connotations embraced by some brands — also communicates sincerity. Green can be active and outdoorsy, and is frequently used by environmental brands to great effect. Yellow communicates competence and happiness, and tends to be bold, much like red. Which dominant color puts the best foot forward in your industry?

For example, when a beer company created a more angular bottle that appeared to have a thicker neck, it was done with the purpose to be perceived as masculine. This helped the company cater to their target audience — and the packaging worked, boosting sales. Shape is important for more than just beer bottling.

These memorable business cards made use of their limited space by changing up their basic shape, such as including the pull-out likeness of the professional on the card itself.A second notable feature of the human genome is the large average gene size of 27, nucleotide pairs.

Every pattern has been test-constructed to verify dimensional accuracy. We say it all the time.


This helped the company cater to their target audience — and the packaging worked, boosting sales. Wiley Encyclopedia of Packaging Technology.

When I was taking package design in college this was my pride and joy and coveted by all.