59
Molecular Organization of Chromosomes Promila Ph.D. Biotechnology GJU S&T Hisar

Molecular organization of chromosomes

Embed Size (px)

Citation preview

Page 1: Molecular organization of chromosomes

Molecular Organization of Chromosomes

PromilaPh.D. BiotechnologyGJU S&T Hisar

Page 2: Molecular organization of chromosomes

Genetic material in a cell

• All cells have the capability to give rise to new cells and the encoded information in a living cell is passed from one generation to another.

•The information encoding material is the genetic or hereditary material of the cell.

Page 3: Molecular organization of chromosomes

Bacterial genetic material

•The prokaryotic (bacterial) genetic material is usually concentrated in a specific clear region of the cytoplasm called nucleiod.

•The bacterial chromosome is a single, circular, double stranded DNA molecule mostly attached to the plasma membrane at one point. It does not contain any histone protein.

• Escherichia coli DNA is circular molecule 4.6 million base pairs in length, containing 4288 annotated protein-coding genes, seven ribosomal RNA (rRNA) operons, and 86 transfer RNA (tRNA) genes. Certain bacteria like the Borrelia burgdorferi possess array of linear chromosome like eukaryotes.

Page 4: Molecular organization of chromosomes
Page 5: Molecular organization of chromosomes

•Besides the chromosomal DNA many bacteria may also carry extra chromosomal genetic elements in the form of small, circular and closed DNA molecules, called plasmids.

• They generally remain floated in the cytoplasm and bear different genes based on which they have been studied.

•Some of the different types of plasmids are F plasmids, R plasmids, virulent plasmids, metabolic plasmids etc.

Page 6: Molecular organization of chromosomes

Virus genetic material

• The chromosomal material of viruses is DNA or RNA which adopts different structures. It is circular when packaged inside the virus particle.

Eukaryotic genetic material

•A Eukaryotic cell has genetic material in the form of genomic DNA enclosed within the nucleus.

• Genes or the hereditary units are located on the chromosomes which exist as chromatin network in the non dividing cell/interphase.

Page 7: Molecular organization of chromosomes

Chromosome

• German biologist Walter Flemming in the early 1880s revealed that during cell division the nuclear material organize themselves into visible thread like structures which were named as chromosomes which stains deep with basic dyes.

•The term chromosome was coined by W. Waldeyer in 1888.

• Chrome is coloured and soma is body, hence they mean “colored bodies” and can be defined as higher order organized arrangement of DNA and proteins.

Page 8: Molecular organization of chromosomes

•It contains many genes or the hereditary units, regulatory elements and other nucleotide sequences.

• Chromosomes also contain DNA-bound proteins, which serve in packaging the DNA and control its functions.

• Chromosomes vary both in number and structure among organisms and the number of chromosomes is characteristic of every species.

•Benden and Bovery in 1887 reported that the number of chromosomes in each species is constant. W.S. Sutton and T. Boveri in 1902 suggested that chromosomes are the physical structures which acted as messengers of heredity.

Page 9: Molecular organization of chromosomes
Page 10: Molecular organization of chromosomes

•Chromosomes are tightly coiled DNA around basic histone proteins, which help in the tight packing of DNA.

• During interphase, the DNA is not tightly coiled into chromosomes, but exists as chromatin.

•In eukaryotes to fit the entire length of DNA in the nucleus it undergoes condensation and the degree to which DNA is condensed is expressed as its packing ratio which is the length of DNA divided by the length into which it is packaged into chromatin along with proteins.

Page 11: Molecular organization of chromosomes

•The shortest human chromosome contains 4.6 x 107 bp of DNA.

• This is equivalent to 14,000 µm of extended DNA.

•In its most condensed state during mitosis, the chromosome is about 2 µm long. This gives a packing ratio of 7000 (14,000/2).

• The DNA is packaged stepwise into the higher order chromatin structure and this is known as “hierarchies of chromosomal organization”.

Page 12: Molecular organization of chromosomes
Page 13: Molecular organization of chromosomes
Page 14: Molecular organization of chromosomes
Page 15: Molecular organization of chromosomes

Structure of Chromosome

•A chromosome at mitotic metaphase consists of two symmetrical structures called chromatids.

• Each chromatid contains a single DNA molecule and both chromatids are attached to each other by centromere and become separated at the beginning of anaphase.

•The chromomeres are bead like accumulations of chromatin material that are sometimes visible along interphase chromosomes.

•The chromomere bearing chromatin has an appearance of a necklace in which several beads occur on a string. Chromomeres are regions of tightly folded DNA and become especially prominent in polytene chromosomes.

Page 16: Molecular organization of chromosomes

Centromere

•Centromere in a chromosome contain specific DNA sequences with special proteins bound to them, forming a disc shaped structure, called kinetochore.

• In electron microscope the kinetochore appears as a plate or cup like disc, 0.20-0.25 nm, in diameter situated upon the primary constriction or centromere.

•The chromosomes of most organisms contain only one centromere and are known as monocentric chromosomes.

Page 17: Molecular organization of chromosomes

•Besides the primary constrictions or centromeres, chromosomes also posses secondary constriction at any point of the chromosome and are constant in their position and extent.

•These constrictions are helpful in identifying particular chromosomes in a set.

• Chromosomes also contain nucleolar organizers which are certain secondary constrictions that contain the genes coding for 5.8S, 18S and 28S ribosomal RNA and induce the formation of nucleoli.

•Sometimes the chromosomes bear round, elongated or knob like appendages known as satellites. The satellite remains connected with the rest of the chromosomes by a thin chromatin filament

Page 18: Molecular organization of chromosomes

Chromatin:

Chemical composition of chromatin Chromatin consists of DNA, RNA and protein. The protein of chromatin could be of two types: histones and non histones.

DNA: •DNA is the most important chemical component of chromatin, since it plays central role of controlling heredity and is most conveniently measured in picograms.

•In addition to describing the genome of an organism by its number of chromosomes, it is also described by the amount of DNA in a haploid cell.

• This is usually expressed as the amount of DNA per haploid cell (usually expressed as picograms) or the number of kilobases per haploid cell and is called the C value.

Page 19: Molecular organization of chromosomes

•This is constant for all cells of a species. For diploid cells it is 2C.

• Extending the C value we reach the C-value paradox.

•One immediate feature of eukaryotic organisms highlights a specific anomaly that was detected early in molecular research. Even though eukaryotic organisms appear to have 2-10 times as many genes as prokaryotes, they have many orders of magnitude more DNA in the cell.

• Furthermore, the amount of DNA per genome is correlated not with the presumed evolutionary complexity of a species. This is stated as the C value paradox: the amount of DNA in the haploid cell of an organism is not related to its evolutionary complexity.

Page 20: Molecular organization of chromosomes

•Lower eukaryotes in general have less DNA, such as nematode Caenorhabditis elegans which has 20 times more DNA than E. coli.

•Vertebrates have greater DNA content about 3pg, in general about 700 times more than E. coli.

•Salamander Amphiuma has a very high DNA content of about 84pg. Man has about 3pg of DNA per haploid genome.

Page 21: Molecular organization of chromosomes

Histones

•Histones are basic proteins as they are enriched with basic proteins arginine and lysine.

•At physiological pH they are cationic and can interact with anionic nucleic acids. They form a highly condensed structure.

•The histones are of five types called H1, H2A H2B, H3, and H4-which are very similar among different species of eukaryotes and have been highly conserved during evolution.

• H1 is the least conserved among all and is also loosely bound with DNA. H1 histone is absent in Sacharomyces cerevisiae.

Page 22: Molecular organization of chromosomes

Non-histones

•In addition to histones the chromatin comprise of many different types of non-histone proteins, which are involved in a range of activities, including DNA replication and gene expression.

•They display more diversity or are not conserved.

•They may also differ between different tissues of same organism.

Page 23: Molecular organization of chromosomes

Euchromatin

•The lightly-stained regions in chromosome when stained with basic dyes are called euchromatin and contain single-copy of genetically-active DNA.

•The extent of chromatin condensation varies during the life cycle of the cell and plays an important role in regulating gene expression.

•In the interphase of cell cycle the chromatin are decondensed and known as euchromatin leading to gene transcription and DNA replication.

Page 24: Molecular organization of chromosomes

Heterochromatin

•The word heterochromatin was coined by Emil Heitz based on cytological observations.

•They are highly condensed and ordered areas in nucleosomal arrays.

•About 10% of interphase chromatin is called heterochromatin and is in a very highly condensed state that resembles the chromatin of cells undergoing mitosis .

•They contain a high density of repetitive DNA found at centromeres and telomeres form heterochromatin.

Page 25: Molecular organization of chromosomes

•Heterochromatin are of two types, the constitutive and facultative heterochromatin.

•The regions that remain condensed throughout the cell cycle are called constitutive heterochromatin whereas the regions where heterochromatin condensation state can change are known as facultative.

• Constitutive heterochromatin is found in the region that flanks the telomeres and centromere of each chromosome and in the distal arm of the Y chromosome in mammals.

•Constitutive heterochromatin possesses very few genes and they also lead to transcriptional inactivation of nearby genes. This phenomenon of gene silencing is known as “position effect”.

Page 26: Molecular organization of chromosomes

•Constitutive heterochromatin also inhibits genetic recombination between homologous repetitive sequences circumventing DNA duplications and deletion.

• Whereas facultative heterochromatin is chromatin that has been specifically inactivated during certain phases of an organism’s life or in certain types of differentiated cells.

•Dosage compensation of X-chromosome or X-chromosome inactivation in mammals is an example of such heterochromatin (Karp 2010).

Page 27: Molecular organization of chromosomes

Centromeres

•Centromeres are those condensed regions within the chromosome that are responsible for the accurate segregation of the replicated chromosome during mitosis and meiosis.

• When chromosomes are stained they typically show a dark-stained region that is the centromere.

• The actual location where the attachments of spindle fibres occur is called the kinetochore and is composed of both DNA and protein.

•The DNA sequence within these regions is called CEN DNA. Because CEN DNA can be moved from one chromosome to another and still provide the chromosome with the ability to segregate, these sequences must not provide any other function.

• Typically CEN DNA is about 120 base pairs long and consists of several sub-domains, CDE-I, CDE-II and CDE-III.

Page 28: Molecular organization of chromosomes
Page 29: Molecular organization of chromosomes

•Mutations in the first two sub-domains have no effect upon segregation, but a point mutation in the CDE-III sub-domain completely eliminates the ability of the centromere to function during chromosome segregation.

•Therefore CDE-III must be actively involved in the binding of the spindle fibers to the centromere.

•The protein component of the kinetochore is only now being characterized. A complex of three proteins called CbfIII binds to normal CDE-III regions but cannot bind to a CDE-III region with a point mutation that prevents mitotic segregation.

•Furthermore, mutants of the genes encoding the Cbf-III proteins also eliminates the ability for chromosomes to segregate during mitosis.

Page 30: Molecular organization of chromosomes

Telomeres

• Telomeres are the region of DNA at the end of the linear eukaryotic chromosome that are required for the replication and stability of the chromosome.

• McClintock recognized their special features when she noticed, that if two chromosomes were broken in a cell, the ends were sticky and end of one could attach to the other and vice versa.

• However she never observed the attachment of the broken end to the end of an unbroken chromosome suggesting that the end of chromosomes have unique features.

Page 31: Molecular organization of chromosomes

•Telomere sequences remain conserved throughout vertebrates and they form caps that protect the chromosomes from nucleases and other destabilizing influences; and they prevent the ends of chromosomes from fusing with one another.

•The telomeric DNA contains direct tandemly repeated sequences of the form (T/A)xGy where x is between 1 and 4 and y is greater than 1. Human telomeres contain the sequence TTAGGG repeated from about 500 to 5000 times.

•Certain bacteria possess telomeres in their linear genetic material which are of two types; one of the types is called a hairpin telomere. As its name implies, the telomeres bend around from the end of one DNA strand to the end of the complimentary strand.

• The other type of telomere is known as an invertron telomere. This type acts to allow an overlap between the ends of the complimentary DNA strands.

Page 32: Molecular organization of chromosomes

Giant chromosomes

• Some cells at certain particular stage of their life cycle contain large nuclei with giant or large sized chromosomes.

•Polytene and lampbrush chromosomes are examples of giant chromosomes.

Page 33: Molecular organization of chromosomes

Polytene Chromosome

•Giant chromosomes were first time observed by E.G. Balbiani in the year 1881 in nuclei of certain secretory cells (salivary glands) of Chironomas larvae (Diptera).

•However he could not conclude them to be chromosomes.

•They were conclusively reported for the first time in insect cells (Drosophila) by Theophilus Painter of the University of Texas in the year 1933.

•Since they were discovered in the salivary glands of insects they were termed as salivary gland chromosomes.

Page 34: Molecular organization of chromosomes

•Cells in the larval salivary gland of Drosophila, mosquito and Chironema contain chromosomes with high DNA content.

• However they may also occur in malphigian tubules, rectum, gut, foot pads, fat bodies, ovarian nurse cells etc.

• Polyteney of giant chromosomes happens by replication of the chromosomal DNA several times without nuclear division (endomitosis) and the resulting daughter chromatids do not separate but remain aligned side by side.

•During endomitosis the nuclear envelope does not rupture and no spindle formation takes place. The polytene chromosomes are visible during interphase and prophase of mitosis.

Page 35: Molecular organization of chromosomes

•They are about 100 times thicker contain 1000 to 2000 chromosomes, than the chromosomes found in most other cells of the organism.

•When stained and viewed under compound microscope at 40X magnification they display about 5000 bands.

•In them the chromomere or the more tightly coiled regions alternate with regions where the DNA fibres are folded loosely.

•A series of dark transverse bands alternates with clear zones of inter bands. Such individual bands can be correlated with particular genes. About 85% of the DNA in polytene chromosomes is in bands and rest 15% is in inter bands.

Page 36: Molecular organization of chromosomes

•The cross banding pattern of each polytene chromosome is a constant characteristic within a species and helps in chromosome mapping during cytogenetic studies.

• In Drosophila melanogaster there are about 5000 bands and 5000 interbands per genome.

• These chromosomes are not inert cellular objects but dynamic structures in which certain regions become “puffed out” due to active DNA transcription at particular stages of development.

Page 37: Molecular organization of chromosomes
Page 38: Molecular organization of chromosomes

•These chromosome puffs are also termed Balbiani rings.

•Puffs may appear and disappear depending on the production of specific proteins which needs to be secreted in large amounts in the larval saliva.

•Another peculiarity of the polytene chromosomes is that the paternal and maternal chromosomes remain associated side by side and the phenomenon is termed somatic pairing.

Page 39: Molecular organization of chromosomes

Lampbrush chromosome

•Lampbrush chromosomes were first observed by Flemming in 1882 in sections of Salamander oocytes and later described by Ruckert in the year 1892.

•They appeared like brushes used for cleaning lamps, hence the name lampbrush chromosome.

•They are transitory structures and can be observed during the diplotene stage of prophase I in meiosis in the oocytes of all animal species both vertebrates and invertebrates.

•They have been described in Sepia (Mollusca), Echinaster (Echinodermata) and in several species of insects, shark, amphibians, reptiles, birds and mammals (humans).

Page 40: Molecular organization of chromosomes

•Lampbrush chromosomes have also been found in spermatocytes of several species, giant nucleus of Acetabularia and even in plants.

• Generally they are smaller in invertebrates than vertebrates.

•They are observed in oocytes because oocytes are high in DNA content.

•Lampbrush chromosomes are functional for studying chromosome organization and genome function during meiotic prophase.

• Additionally lampbrush chromosomes are widely used for construction of detail cytological maps of individual chromosomes.

Page 41: Molecular organization of chromosomes
Page 42: Molecular organization of chromosomes

•They are of exceptionally large sizes and present in bivalent form.

• They are formed due to the active synthesis of mRNA molecules for future use by the egg cells, when no synthesis of mRNA molecule is possible during the mitotic cell division.

•Lampbrush chromosomes are clearly visible in the light microscope they are organized into a series of chromomeres with large chromatin symmetrical loops extending laterally.

Page 43: Molecular organization of chromosomes

•Each loop appears at a constant position in the chromosome (10,000 loops per chromosome set or haploid set).

•Each loop has an axis made up of DNA unfolded from the chromosome and is transcriptionally highly active.

•Wherein several transcription units with polarized RNP-matrix coats the DNA axis of the loop.

• The majority of the DNA, however, is not in loops but remains highly condensed in the chromomeres on the axis and lacks expression of genes.

Page 44: Molecular organization of chromosomes

•The loops perform intense transcription of heterogenous RNA (precursors of mRNA molecules for ribosomal and histone proteins).

•Thus each lateral loop is covered by an assymetrical matrix of RNA transcripts; thicker at one end of the loop than other.

•The number of pairs of loops gradually increases during meiosis till it reaches maximum at diplotene.

•This stage may persist for months or years as oocytes build up supply of mRNA required for further development. As meiosis proceeds further number of loops gradually decrease and loops ultimately disappear due to reabsorption into the chromosome or disintegration.

Page 45: Molecular organization of chromosomes

Genome Complexity

•Forty years ago it was thought that the amount of DNA in a genome correlated with the complexity of an organism.

•Back then, you often saw graphs like the one on the botttom. The idea was that the more complex the species the more genes it needed. Preliminary data seemed to confirm this idea.

Page 46: Molecular organization of chromosomes

•In the late 1960's scientists started looking at the complexity of the genome itself.

• They soon discovered that large genomes were often composed of huge amounts of repetitive sequences.

•The amount of "unique sequence" DNA was only a few percent of the total DNA in these large genomes.

•This gave rise to the concept of junk DNA and the recognition that genome size was not a reliable indicator of the number of genes.

•Today we know that there isn't a direct correlation between genome size and complexity.

Page 47: Molecular organization of chromosomes

• Recent data, reveals that the range of DNA sizes in many groups can vary over several orders of magnitude.

• Mammals don't have any more DNA in their genome than most flowering plants (angiosperms). Or even gymnosperms, for that matter.

•So complexity can be "broadly defined as the number and different types of cells, and the degree of cellular organization."

•The genomes of most eukaryotes are larger and more complex than those of prokaryotes. This larger size of eukaryotic genomes is not inherently surprising, since one would expect to find more genes in organisms that are more complex.

•However, the genome size of many eukaryotes does not appear to be related to genetic complexity. For example, the genomes of salamanders and lilies contain more than ten times the amount of DNA that is in the human genome, yet these organisms are clearly not ten times more complex than humans.

Page 48: Molecular organization of chromosomes

Fig. Genome size. The range of sizes of the genomes of representative groups of organisms are shown on a logarithmic scale

Page 49: Molecular organization of chromosomes

•This apparent paradox was resolved by the discovery that the genomes of most eukaryotic cells contain not only functional genes but also large amounts of DNA sequences that do not code for proteins.

• The difference in the sizes of the salamander and human genomes thus reflects larger amounts of non-coding DNA, rather than more genes, in the genome of the salamander.

•The presence of large amounts of non-coding sequences is a general property of the genomes of complex eukaryotes. Thus, the thousand fold greater size of the human genome compared to that of E. coli is not due solely to a larger number of human genes.

• The human genome is thought to contain approximately 100,000 genes—only about 25 times more than E. coli has. Much of the complexity of eukaryotic genomes thus results from the abundance of several different types of noncoding sequences, which constitute most of the DNA of higher eukaryotic cells.

Page 50: Molecular organization of chromosomes

The structure of eukaryotic genes

•Most eukaryotic genes contain segments of coding sequences (exons) interrupted by noncoding sequences (introns). Both exons and introns are transcribed to yield a long primary RNA transcript. The introns are then removed by splicing to form the mature mRNA.

•Introns were first discovered in 1977, independently in the laboratories of Phillip Sharp and Richard Roberts, during studies of the replication of adenovirus in cultured human cells.

Page 51: Molecular organization of chromosomes

•Soon after the discovery of introns in adenovirus, similar observations were made on cloned genes of eukaryotic cells.

• For example, electron microscopic analysis of RNA-DNA hybrids and subsequent nucleotide sequencing of cloned genomic DNAs and cDNAs indicated that the coding region of the mouse β-globin gene (which encodes the β subunit of hemoglobin) is interrupted by two introns that are removed from the mRNA by splicing.

•The intron-exon structure of many eukaryotic genes is quite complicated, and the amount of DNA in the intron sequences is often greater than that in the exons

Page 52: Molecular organization of chromosomes

•Most introns have no known cellular function, although a few have been found to encode functional RNAs or proteins.

• Introns are generally thought to represent remnants of sequences that were important earlier in evolution.

•In particular, introns may have helped accelerate evolution by facilitating recombination between protein-coding regions (exons) of different genes—a process known as exon shuffling.

•Exons frequently encode functionally distinct protein domains, so recombination between introns of different genes would result in new genes containing novel combinations of protein-coding sequences.

• As predicted by this hypothesis, DNA sequencing studies have demonstrated that some genes are chimeras of exons derived from several other genes, providing direct evidence that new genes can be formed by recombination between intron sequences.

Page 53: Molecular organization of chromosomes

•It appears most likely that introns were present early in evolution, prior to the divergence of prokaryotic and eukaryotic cells.

•According to this hypothesis, introns played an important role in the initial assembly of protein-coding sequences in the ancient ancestors of present-day cells.

• Introns were subsequently lost from most genes of prokaryotes and simpler eukaryotes (e.g., yeasts) in response to evolutionary selection for rapid replication, which led to streamlining the genomes of these organisms.

•However, since rapid cell division is not an advantage to higher eukaryotes, introns have been retained in their genomes.

• Alternatively, introns may have arisen later in evolution as a result of the insertion of DNA sequences into genes that had already been formed as continuous protein-coding sequences

Page 54: Molecular organization of chromosomes

Gene Families and Pseudogenes

•Another factor contributing to the large size of eukaryotic genomes is that some genes are repeated many times.

•Whereas most prokaryotic genes are represented only once in the genome, many eukaryotic genes are present in multiple copies, called gene families.

•In some cases, multiple copies of genes are needed to produce RNAs or proteins required in large quantities, such as ribosomal RNAs or histones.

• In other cases, distinct members of a gene family may be transcribed in different tissues or at different stages of development.

•For example, the α and β subunits of hemoglobin are both encoded by gene families in the human genome, with different members of these families being expressed in embryonic, fetal, and adult tissues.

Page 55: Molecular organization of chromosomes

•Gene families are thought to have arisen by duplication of an original ancestral gene, with different members of the family then diverging as a consequence of mutations during evolution.

•As might be expected, however, not all mutations enhance gene function.

• Some gene copies have instead sustained mutations that result in their loss of ability to produce a functional gene product.

• For example, the human α- and β-globin gene families each contain two genes that have been inactivated by mutations.

•Such nonfunctional gene copies (called pseudogenes) represent evolutionary relics that significantly increase the size of eukaryotic genomes without making a functional genetic contribution.

Page 56: Molecular organization of chromosomes

Repetitive DNA Sequences

•A substantial portion of eukaryotic genomes consists of highly repeated non- coding DNA sequences.

• These sequences, sometimes present in hundreds of thousands of copies per genome, were first demonstrated by Roy Britten and David Kohne during studies of the rates of reassociation of denatured fragments of cellular DNAs.

•Experiments indicated that approximately 40% of mammalian DNA consists of highly repetitive sequences, some of which are repeated 105 to 106 times.

•Further analysis has identified several types of these highly repeated sequences.

Page 57: Molecular organization of chromosomes

•One class (called simple-sequence DNA) contains tandem arrays of thousands of copies of short sequences, ranging from 5 to 200 nucleotides.

• For example, one type of simple-sequence DNA in Drosophila consists of tandem repeats of the seven nucleotide unit ACAAACT.

•Because of their distinct base compositions, many simple-sequence DNAs can be separated from the rest of the genomic DNA by equilibrium centrifugation in CsCl density gradients.

•Since such repeat-sequence DNAs band as “satellites” separate from the main band of DNA, they are frequently referred to as satellite DNAs.

•These sequences are repeated millions of times per genome, accounting for 10 to 20% of the DNA of most higher eukaryotes.

•Simple-sequence DNAs are not transcribed and do not convey functional genetic information.

Page 58: Molecular organization of chromosomes

The Number of Genes in Eukaryotic Cells

•In bacterial genomes, most of the DNA encodes proteins. For example, the genome of E. coli is approximately 4.6 × 106 base pairs long and contains 4288 genes, with nearly 90% of the DNA used as protein-coding sequence.

•The genomes of higher animals (such as humans) are still more complex and contain large amounts of noncoding DNA. Thus, only a small fraction of the 3 × 109 base pairs of the human genome is expected to correspond to protein-coding sequence. Approximately one-third of the genome corresponds to highly repetitive sequences, leaving an estimated 2 × 109 base pairs for functional genes, pseudogenes, and nonrepetitive spacer sequences.

Organism Genome size (Mb)

Protein-coding

sequence (percent)

Number of genes

E. coli 4.6 90 4,288

S. cerevisiae 12 70 5,885

C. elegans 97 25 19,099

Drosophila 180 13 13,600

Human 3,000 3 100,000

Page 59: Molecular organization of chromosomes

Thank You