Eukaryotic chromosome structure
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Eukaryotic chromosome structure refers to the levels of packaging from raw DNA molecules to the
chromosomal A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
structures seen during
metaphase Metaphase ( and ) is a stage of mitosis in the eukaryotic cell cycle in which chromosomes are at their second-most condensed and coiled stage (they are at their most condensed in anaphase). These chromosomes, carrying genetic information, align ...
in
mitosis In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
or
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
. Chromosomes contain long strands of DNA containing genetic information. Compared to prokaryotic chromosomes,
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
chromosomes are much larger in size and are linear chromosomes. Eukaryotic chromosomes are also stored in the
cell nucleus The cell nucleus (pl. nuclei; from Latin or , meaning ''kernel'' or ''seed'') is a membrane-bound organelle found in eukaryotic cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, h ...
, while chromosomes of prokaryotic cells are not stored in a nucleus. Eukaryotic chromosomes require a higher level of packaging to condense the DNA molecules into the cell nucleus because of the larger amount of DNA. This level of packaging includes the wrapping of DNA around proteins called histones in order to form condensed nucleosomes.


History

The double helix was discovered in 1953 by
James Watson James Dewey Watson (born April 6, 1928) is an American molecular biologist, geneticist, and zoologist. In 1953, he co-authored with Francis Crick the academic paper proposing the double helix structure of the DNA molecule. Watson, Crick and ...
and
Francis Crick Francis Harry Compton Crick (8 June 1916 – 28 July 2004) was an English molecular biologist, biophysicist, and neuroscientist. He, James Watson, Rosalind Franklin, and Maurice Wilkins played crucial roles in deciphering the helical struc ...
. Other researchers made very important, but unconnected findings about the composition of DNA. Ultimately it was
Watson and Crick "Molecular Structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid" was the first article published to describe the discovery of the double helix structure of DNA, using X-ray diffraction and the mathematics of a helix transform. ...
who put all of these findings together to come up with a model for DNA. Later, chemist Alexander Todd determined that the backbone of a DNA molecule contained repeating phosphate and deoxyribose sugar groups. The biochemist
Erwin Chargaff Erwin Chargaff (11 August 1905 – 20 June 2002) was an Austro-Hungarian-born American biochemist, writer, Bucovinian Jew who emigrated to the United States during the Nazi Germany, Nazi era, and professor of biochemistry at Columbia University ...
found that
adenine Adenine () ( symbol A or Ade) is a nucleobase (a purine derivative). It is one of the four nucleobases in the nucleic acid of DNA that are represented by the letters G–C–A–T. The three others are guanine, cytosine and thymine. Its derivati ...
and
thymine Thymine () ( symbol T or Thy) is one of the four nucleobases in the nucleic acid of DNA that are represented by the letters G–C–A–T. The others are adenine, guanine, and cytosine. Thymine is also known as 5-methyluracil, a pyrimidine nu ...
always paired while
cytosine Cytosine () ( symbol C or Cyt) is one of the four nucleobases found in DNA and RNA, along with adenine, guanine, and thymine (uracil in RNA). It is a pyrimidine derivative, with a heterocyclic aromatic ring and two substituents attached (an am ...
and
guanine Guanine () ( symbol G or Gua) is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine (uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside is called ...
always paired. High resolution X-ray images of DNA that were obtained by
Maurice Wilkins Maurice Hugh Frederick Wilkins (15 December 1916 – 5 October 2004) was a New Zealand-born British biophysicist and Nobel laureate whose research spanned multiple areas of physics and biophysics, contributing to the scientific understanding o ...
and
Rosalind Franklin Rosalind Elsie Franklin (25 July 192016 April 1958) was a British chemist and X-ray crystallographer whose work was central to the understanding of the molecular structures of DNA (deoxyribonucleic acid), RNA (ribonucleic acid), viruses, co ...
suggested a helical, or corkscrew like shape. Some of the first scientists to recognize the structures now known as chromosomes were
Schleiden Schleiden is a town in North Rhine-Westphalia, Germany. It lies in the Eifel hills, in the district of Euskirchen, and has 12,998 inhabitants as of 30 June 2017. Schleiden is connected by a tourist railway to Kall, on the Eifel Railway between Col ...
,
Virchow Rudolf Ludwig Carl Virchow (; or ; 13 October 18215 September 1902) was a German physician, anthropologist, pathologist, prehistorian, biologist, writer, editor, and politician. He is known as "the father of modern pathology" and as the founder ...
, and Bütschli. The term was coined by Heinrich Wilhelm Gottfried von Waldeyer-Hartz, referring to the term
chromatin Chromatin is a complex of DNA and protein found in eukaryotic cells. The primary function is to package long DNA molecules into more compact, denser structures. This prevents the strands from becoming tangled and also plays important roles in r ...
, was introduced by
Walther Flemming Walther Flemming (21 April 1843 – 4 August 1905) was a German biologist and a founder of cytogenetics. He was born in Sachsenberg (now part of Schwerin) as the fifth child and only son of the psychiatrist Carl Friedrich Flemming (1799–1880) ...
. Scientists also discovered plant and animal cells have a central compartment called the
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucle ...
. They soon realized chromosomes were found inside the nucleus and contained different information for many different traits.


Structure

In eukaryotes, such as humans, roughly 3.2 billion
nucleotide Nucleotides are organic molecules consisting of a nucleoside and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both of which are essential biomolecules wi ...
s are spread out over 23 different chromosomes (males have both an
X chromosome The X chromosome is one of the two sex-determining chromosomes (allosomes) in many organisms, including mammals (the other is the Y chromosome), and is found in both males and females. It is a part of the XY sex-determination system and XO sex-d ...
and a
Y chromosome The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abse ...
instead of a pair of X chromosomes as seen in females). Each chromosome consists enormously long linear DNA molecule associated with proteins that fold and pack the fine thread of DNA into a more compact structure. Commonly, many people think the structure of a chromosome is in an "X" shape. But this is only present when the cell divides. Researchers have now been able to model the structure of chromosomes when they are active. This is extremely important because the way that DNA folds up in chromosome structures is linked to the way DNA is used. Scientists have been able to develop the 3D structures of chromosomes in a single cell. The scientists used hundreds of measurements of where different parts of the DNA get close to one another to help create this model. This research was done by scientists at the Department of Biochemistry at Cambridge, working with others from the Babraham Institute and the
Weizmann Institute The Weizmann Institute of Science ( he, מכון ויצמן למדע ''Machon Vaitzman LeMada'') is a public research university in Rehovot, Israel, established in 1934, 14 years before the State of Israel. It differs from other Israeli univ ...
.


Nucleosomes

The
nucleosome A nucleosome is the basic structural unit of DNA packaging in eukaryotes. The structure of a nucleosome consists of a segment of DNA wound around eight histone proteins and resembles thread wrapped around a spool. The nucleosome is the fundamen ...
is the basic unit of DNA condensation and consists of a DNA double helix bound to an octamer of core
histones In biology, histones are highly basic proteins abundant in lysine and arginine residues that are found in eukaryotic cell nuclei. They act as spools around which DNA winds to create structural units called nucleosomes. Nucleosomes in turn are wr ...
(2 dimers of H2A and H2B, and an H3/H4
tetramer A tetramer () (''tetra-'', "four" + '' -mer'', "parts") is an oligomer formed from four monomers or subunits. The associated property is called ''tetramery''. An example from inorganic chemistry is titanium methoxide with the empirical formula Ti ...
). About 147
base pairs A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
of DNA coil around 1 octamer, and ~20 base pairs are sequestered by the addition of the linker histone (H1), and various length of "linker" DNA (~0-100 bp) separate the nucleosomes. The spacing of nucleosomes along DNA results in a “beads on a string” appearance. Histone modification controls the accessibility to DNA. Histone acetyltransferases or HATs acetylate residues on the histone tail leading to increased accessibility to DNA.


Packaging

Packaging of DNA is facilitated by the electrostatic charge distribution:
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phospho ...
groups cause DNA to have a negative charge, whilst the histones are positively charged. Most eukaryotic cells contain histones (with a few exceptions) as well as the kingdom
Archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
. Specifically histones H3 and H4 are nearly identical in structure among all eukaryotes, suggesting strict evolutionary conservation of both structure and function. Histones are positively charged molecules as they contain
lysine Lysine (symbol Lys or K) is an α-amino acid that is a precursor to many proteins. It contains an α-amino group (which is in the protonated form under biological conditions), an α-carboxylic acid group (which is in the deprotonated −C ...
and
arginine Arginine is the amino acid with the formula (H2N)(HN)CN(H)(CH2)3CH(NH2)CO2H. The molecule features a guanidino group appended to a standard amino acid framework. At physiological pH, the carboxylic acid is deprotonated (−CO2−) and both the am ...
in larger quantities and DNA is negatively charged. This allows histones to make a strong ionic bond to DNA form a nucleosome. The most basic level of DNA condensation is the wrapping of DNA around the histone core proteins. Higher-order packaging is accomplished by specialized proteins that bind and fold the DNA. This generates a series of loops and coils that provide increasingly higher levels of organization and prevent the DNA from becoming tangled and unmanageable. This complex of DNA and proteins are called chromatin. But in addition to proteins involved with packaging, chromosomes are associated with proteins involved with
DNA replication In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all living organisms acting as the most essential part for biological inheritanc ...
,
DNA repair DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA dam ...
, and
gene expression Gene expression is the process by which information from a gene is used in the synthesis of a functional gene product that enables it to produce end products, protein or non-coding RNA, and ultimately affect a phenotype, as the final effect. The ...
.


References

{{reflist Molecular genetics DNA