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Chromosome-upright

A scheme of a condensed (metaphase) chromosome. (1) Chromatid - one of the two identical parts of the chromosome after S phase. (2) Centromere - the point where the two chromatids touch, and where the microtubules attach. (3) Short arm. (4) Long arm.

Chromosomes are organized structures of DNA and proteins that are found in cells. Chromosomes contain a single continuous piece of DNA, which contains many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions. The word chromosome comes from the Greek χρῶμα (chroma, color) and σῶμα (soma, body) due to their property of being stained very strongly by some dyes.

Chromosomes vary extensively between different organisms. The DNA molecule may be circular or linear, and can contain anything from tens of kilobase pairs to hundreds of megabase pairs. Typically eukaryotic cells (cells with nuclei) have large linear chromosomes and prokaryotic cells (cells without nuclei) smaller circular chromosomes, although there are many exceptions to this rule. Furthermore, cells may contain more than one type of chromosome; for example mitochondria in most eukaryotes and chloroplasts in plants have their own small chromosomes.

In eukaryotes, nuclear chromosomes are packaged by proteins into a condensed structure called chromatin. This allows the massively-long DNA molecules to fit into the cell nucleus. The structure of chromatin varies through the cell cycle, and is responsible for the organisation of chromosomes into the classic four-arm structure during mitosis and meiosis.

"Chromosome" is a rather loosely defined term. In prokaryotes, a small circular DNA molecule may be called either a plasmid or a small chromosome. These small circular genomes are also found in mitochondria and chloroplasts, reflecting their bacterial origins. The simplest chromosomes are found in viruses: these DNA or RNA molecules are short linear or circular chromosomes that often lack any structural proteins.

History

This is a brief history of research in a complex field where each advance was hard won, and often hotly disputed at the time.

Visual discovery of chromosomes. Textbooks have often said that chromosomes were first observed in plant cells by a Swiss botanist named Karl Wilhelm von Nägeli in 1842.[1] However, this opinion has been challenged, perhaps decisively, by Henry Harris, who has freshly reviewed the primary literature.[2] In his opinion the claim of Nägeli to have seen spore mother cells divide is mistaken, as are some of his interpretations. Harris considers other candidates, especially Wilhelm Hofmeister, whose publications in 1848-9 include plates which definitely show mitotic events.[3][4] Hofmeister was also the choice of Cyril Darlington.

The work of other cytologists such as Walther Flemming, Eduard Strasburger, Otto Bütschli, Oskar Hertwig and Carl Rabl should definitely be acknowledged. The use of basophilic aniline dyes was a new technique for effectively staining the chromatin material in the nucleus. Their behavior in animal (salamander) cells was later described in detail by Walther Flemming, who in 1882 "provided a superb summary of the state of the field".[5][6] The name chromosome was invented in 1888 by Heinrich von Waldeyer. However, van Beneden's monograph of 1883 on the fertilised eggs of the parasitic roundworm Ascaris maglocephala was the outstanding work of this period.[7] His conclusions are classic:

  • Thus there is no fusion between the male chromatin and the female chromatin at any stage of division...
  • The elements of male origin and those of female origin are never fused together in a cleavage nucleus, and perhaps they remain distinct in all the nuclei derived from them. [tranl: Harris p162] chat alot of uterous

"It is not easy to identify who first discerned chromosomes during mitosis, but there is no doubt that those who first saw them had no idea of their significance... [but] with the work of Balbiani and van Beneden we move away from... the mechanism of cell division to a precise delineation of chromosomes and what they do during the division of the cell." [8]

Van Beneden's master work was closely followed by that of Carl Rabl, who reached similar conclusions. [9] This more or less concludes the first period, in which chromosomes were visually sighted, and the morphological stages of mitosis were described. Coleman also gives a useful review of these discoveries.[10]

Nucleus as the seat of heredity. The origin of this epoch-making idea lies in a few sentences tucked away in Ernst Haeckel's Generelle Morphologie of 1866.[11] The evidence for this insight gradually acumulated until, after twenty or so years, two of the greatest in a line of great German scientists spelt it out. August Weismann proposed that the germ line was separate from the soma, and that the cell nucleus was the repository of the hereditary material, which he proposed was arranged along the chromosomes in a linear manner. Furthermore, he proposed that at fertilisation a new ombination of chromosomes (and their hereditary material) would be formed. This was the explanation for the reduction division of meiosis (first described by van Beneden).

Chromosomes as vectors of heredity. In a series of outstanding experiments, Theodor Boveri gave the definitive demonstration that chromosomes were the vectors of heredity. His two principles were:

The continuity of chromosomes
The individuality of chromosomes.

It was the second of these principles which was so original. He was able to test the proposal put forward by Wilhelm Roux, that each chromosome carries a different genetic load, and showed that Roux was right. Upon the rediscovery of Mendel, Boveri was able to point out the connection between the rules of inheritance and the behaviour of the chromosomes. It is interesting to see that Boveri influenced two generations of American cytologists: Edmund Beecher Wilson, Walter Sutton and Theophilus Painter were all influenced by Boveri (Wilson and Painter actually worked with him). In his famous textbook The Cell, Wilson linked Boveri and Sutton together by the Boveri-Sutton theory. Mayr remarks that the theory was hotly contested by some famous geneticists: William Bateson, Wilhelm Johannsen, Richard Goldschmidt and T.H. Morgan, all of a rather dogmatic turn of mind. Eventually complete proof came from chromosome maps – in Morgan's own lab! [12]

Chromosomes in eukaryotes

Mergefrom
It has been suggested that Eukaryotic chromosome fine structure be merged into this article or section. (Discuss)
Eukaryotes (cells with nuclei such as plants, yeast, and animals) possess multiple large linear chromosomes contained in the cell's nucleus. Each chromosome has one centromere, with one or two arms projecting from the centromere, although under most circumstances these arms are not visible as such. In addition most eukaryotes have a small circular mitochondrial genome, and some eukaryotes may have additional small circular or linear cytoplasmic chromosomes.

In the nuclear chromosomes of eukaryotes, the uncondensed DNA exists in a semi-ordered structure, where it is wrapped around histones (structural proteins), forming a composite material called chromatin.

Chromatin

Main article: Chromatin
Chromatin Structures

Fig. 2: The major structures in DNA compaction; DNA, the nucleosome, the 10nm "beads-on-a-string" fibre, the 30nm fibre and the metaphase chromosome.

Chromatin is the complex of DNA and protein found in the eukaryotic nucleus which packages chromosomes. The structure of chromatin varies significantly between different stages of the cell cycle, according to the requirements of the DNA.

Interphase chromatin

During interphase (the period of the cell cycle where the cell is not dividing) two types of chromatin can be distinguished:

  • Euchromatin, which consists of DNA that is active, e.g., expressed as protein.
  • Heterochromatin, which consists of mostly inactive DNA. It seems to serve structural purposes during the chromosomal stages. Heterochromatin can be further distinguished into two types:
    • Constitutive heterochromatin, which is never expressed. It is located around the centromere and usually contains repetitive sequences.
    • Facultative heterochromatin, which is sometimes expressed.

Individual chromosomes cannot be distinguished at this stage - they appear in the nucleus as a homogeneous tangled mix of DNA and protein.

Metaphase chromatin and division

See also: mitosis and meiosis
File:HumanChromosomesChromomycinA3.jpg

In the early stages of mitosis or meiosis (cell division), the chromatin strands become more and more condensed. They cease to function as accessible genetic material (transcription stops) and become a compact transportable form. This compact form makes the individual chromosomes visible, and they form the classic four arm structure, a pair of sister chromatids attached to each other at the centromere. The shorter arms are called p arms (from the French petit, small) and the longer arms are called q arms (q follows p in the Latin alphabet). This is the only natural context in which individual chromosomes are visible with an optical microscope.

During divisions long microtubules attach to the centromere and the two opposite ends of the cell. The microtubules then pull the chromatids apart, so that each daughter cell inherits one set of chromatids. Once the cells have divided, the chromatids are uncoiled and can function again as chromatin. In spite of their appearance, chromosomes are structurally highly condensed which enables these giant DNA structures to be contained within a cell nucleus (Fig. 2).

The self assembled microtubules form the spindle, which attaches to chromosomes at specialized structures called kinetochores, one of which is present on each sister chromatid. A special DNA base sequence in the region of the kinetochores provides, along with special proteins, longer-lasting attachment in this region.

Chromosomes in prokaryotes

The prokaryotes - bacteria and archaea - typically have a single circular chromosome, but many variations do exist.[13] Most bacteria have a single circular chromosome that can range in size from only 160,000 base pairs in the endosymbiotic bacteria Candidatus Carsonella ruddii,[14] to 12,200,000 base pairs in the soil-dwelling bacteria Sorangium cellulosum.[15] Spirochaetes of the genus Borrelia are a notable exception to this arrangement, with bacteria such as Borrelia burgdorferi, the cause of Lyme disease, containing a single linear chromosome.[16]

Structure in sequences

Prokaryotes chromosomes have less sequence-based structure than eukaryotes. Bacteria typically have a single point (the origin of replication) from which replication starts, while some archaea contain multiple replication origins.[17] The genes in prokaryotes are often organised in operons, and do not contain introns, unlike eukaryotes.

DNA packaging

Prokaryotes do not possess nuclei, instead their DNA is organized into a structure called the nucleoid.[18] The nucleoid is a distinct structure and occupies a defined region of the bacterial cell. This structure is however dynamic and is maintained and remodeled by the actions of a range of histone-like proteins, which associate with the bacterial chromosome.[19] In archaea, the DNA in chromosomes is even more organized, with the DNA packaged within structures similar to eukaryotic nucleosomes.[20][21]

Bacterial chromosomes tend to be tethered to the plasma membrane of the bacteria. In molecular biology application, this allows for its isolation from plasmid DNA by centrifugation of lysed bacteria and pelleting of the membranes (and the attached DNA).

Prokaryotic chromosomes and plasmids are, like eukaryotic DNA, generally supercoiled. The DNA must first be released into its relaxed state for access for transcription, regulation, and replication.

Number of chromosomes in various organisms

Eukaryotes

Chromosome numbers (2n) in some plants
Plant Species #
Arabidopsis thaliana 10
Rye 14
Maize 20
Einkorn wheat[22] 14
Durum wheat[22] 28
Bread wheat[22] 42
Wild tobacco[How to reference and link to summary or text] 24
Cultivated tobacco 48
Adder's Tongue Fern[23] 1262
Chromosome numbers (2n) in some animals
Species # Species #
Common fruit fly 8 Guinea Pig [24] 64
Dove[How to reference and link to summary or text] 16 Snail[How to reference and link to summary or text] 24
Earthworm[25] 36 Tibetan fox 36
Domestic cat 38 Domestic pig 38
Lab mouse 40 Lab rat 42
Rabbit[How to reference and link to summary or text] 44 Syrian hamster 44
Hare[How to reference and link to summary or text] 46 Human[26] 46
Gorillas, Chimpanzees[26] 48 Domestic sheep 54
Elephants[27] 56 Cow 60
Donkey 62 Horse 64
Dog[28] 78 Chicken[29] 39
Goldfish[30] 100-104 Silkworm[31] 28
Chromosome numbers in other organisms
Species Large
Chromosomes
Intermediate
Chromosomes
Small
Chromosomes
Trypanosoma brucei 11 6 ~100
File:PLoSBiol3.5.Fig1bNucleus46Chromosomes.jpg

Normal members of a particular eukaryotic species all have the same number of nuclear chromosomes (see the table). Other eukaryotic chromosomes, i.e., mitochondrial and plasmid-like small chromosomes, are much more variable in number, and there may be thousands of copies per cell.

Asexually reproducing species have one set of chromosomes, which is the same in all body cells.

Sexually reproducing species have somatic cells (body cells), which are diploid [2n] having two sets of chromosomes, one from the mother and one from the father. Gametes, reproductive cells, are haploid [n]: they have one set of chromosomes. Gametes are produced by meiosis of a diploid germ line cell. During meiosis, the matching chromosomes of father and mother can exchange small parts of themselves (crossover), and thus create new chromosomes that are not inherited solely from either parent. When a male and a female gamete merge (fertilization), a new diploid organism is formed.

Some animal and plant species are polyploid [Xn]: they have more than two sets of homologous chromosomes. Agriculturally important plants such as tobacco or wheat are often polyploid compared to their ancestral species. Wheat has a haploid number of seven chromosomes, still seen in some cultivars as well as the wild progenitors. The more common pasta and bread wheats are polyploid, having 28 (tetraploid) and 42 (hexaploid) chromosomes compared to the 14 (diploid) chromosomes in the wild wheat.[32]

Prokaryotes

Prokaryote species generally have one copy of each major chromosome, but most cells can easily survive with multiple copies.[33] Plasmids and plasmid-like small chromosomes are, like in eukaryotes, very variable in copy number. The number of plasmids in the cell is almost entirely determined by the rate of division of the plasmid - fast division causes high copy number, and vice versa.

Karyotype

Main article: Karyotype
NHGRI human male karyotype

Figure 3: Karyogram of a human male

In general, the karyotype is the characteristic chromosome complement of a eukaryote species.[34] The preparation and study of karyotypes is part of cytogenetics.

Although the replication and transcription of DNA is highly standardized in eukaryotes, the same cannot be said for their karotypes, which are often highly variable. There may be variation between species in chromosome number and in detailed organization. In some cases there is significant variation within species. Often there is variation 1. between the two sexes. 2. between the germ-line and soma (between gametes and the rest of the body). 3. between members of a population, due to balanced genetic polymorphism. 4. geographical variation between races. 5. mosaics or otherwise abnormal individuals. Finally, variation in karyotype may occur during development from the fertilised egg.

The technique of determining the karyotype is usually called karyotyping. Cells can be locked part-way through division (in metaphase) in vitro (in a reaction vial) with colchicine. These cells are then stained, photographed and arranged into a karyogram, with the set of chromosomes arranged, autosomes in order of length, and sex chromosomes (here XY) at the end: Fig. 3.

Like many sexually reproducing species, humans have special gonosomes (sex chromosomes, in contrast to autosomes). These are XX in females and XY in males.

Historical note

Investigation into the human karyotype took many years to settle the most basic question: how many chromosomes does a normal diploid human cell contain? In 1912, Hans von Winiwarter reported 47 chromosomes in spermatogonia and 48 in oogonia, concluding an XX/XO sex determination mechanism.[35] Painter in 1922 was not certain whether the diploid number of man was 46 or 48, at first favouring 46.[36] He revised his opinion later from 46 to 48, and he correctly insisted on man having an XX/XY system.[37] Considering their techniques, these results were quite remarkable.

New techniques were needed to definitively solve the problem:

1. Using cells in culture
2. Pretreating cells in a hypotonic solution, which swells them and spreads the chromosomes
3. Arresting mitosis in metaphase by a solution of colchicine
4. Squashing the preparation on the slide forcing the chromosomes into a single plane
5. Cutting up a photomicrograph and arranging the result into an indisputable karyogram.

It took until the mid 1950s until it became generally accepted that the karyotype of man included only 46 chromosomes.[38][39] Rather interestingly, chimpanzees (our closest living relatives) have 48 chromosomes.

Chromosomal aberrations

Main article: Chromosome abnormalities
Single Chromosome Mutations

The three major single chromosome mutations; deletion (1), duplication (2) and inversion (3).

Two Chromosome Mutations

The two major two-chromosome mutations; insertion (1) and translocation (2).

Chromosome 21

In Down syndrome, there are three copies of chromosome 21

Chromosomal aberrations are disruptions in the normal chromosomal content of a cell, and are a major cause of genetic conditions in humans, such as Down syndrome. Some chromosome abnormalities do not cause disease in carriers, such as translocations, or chromosomal inversions, although they may lead to a higher chance of having a child with a chromosome disorder. Abnormal numbers of chromosomes or chromosome sets, aneuploidy, may be lethal or give rise to genetic disorders. Genetic counseling is offered for families that may carry a chromosome rearrangement.

Chromosomal mutations produce changes in whole chromosomes (more than one gene) or in the number of chromosomes present.

  • Deletion - loss of part of a chromosome
  • Duplication - extra copies of a part of a chromosome
  • Inversion - reverse the direction of a part of a chromosome
  • Translocation - part of a chromosome breaks off and attaches to another chromosome

Most mutations are neutral - have little or no effect

A detailed graphical display of all human chromosomes and the diseases annotated at the correct spot may be found at [1].

Human chromosomes

Human cells have 23 pairs of large linear nuclear chromosomes, giving a total of 46 per cell. In addition to these, human cells have many hundreds of copies of the mitochondrial genome. Sequencing of the human genome has provided a great deal of information about each of the chromosomes. Below is a table compiling statistics for the chromosomes, based on the Sanger Institute's human genome information in the Vertebrate Genome Annotation (VEGA) database.[40] Number of genes is an estimate as it is in part based on gene predictions. Total chromosome length is an estimate as well, based on the estimated size of unsequenced heterochromatin regions.

Chromosome Genes Total bases Sequenced bases[41]
1 3,148 247,200,000 224,999,719
2 902 242,750,000 237,712,649
3 1,436 199,450,000 194,704,827
4 453 191,260,000 187,297,063
5 609 180,840,000 177,702,766
6 1,585 170,900,000 167,273,992
7 1,824 158,820,000 154,952,424
8 781 146,270,000 142,612,826
9 1,229 140,440,000 120,312,298
10 1,312 135,370,000 131,624,737
11 405 134,450,000 131,130,853
12 1,330 132,290,000 130,303,534
13 623 114,130,000 95,559,980
14 886 106,360,000 88,290,585
15 676 100,340,000 81,341,915
16 898 88,820,000 78,884,754
17 1,367 78,650,000 77,800,220
18 365 76,120,000 74,656,155
19 1,553 63,810,000 55,785,651
20 816 62,440,000 59,505,254
21 446 46,940,000 34,171,998
22 595 49,530,000 34,893,953
X (sex chromosome) 1,093 154,910,000 151,058,754
Y (sex chromosome) 125 57,740,000 22,429,293

See also

External links

References

  1. Nägeli C. 1842. Zur Entwickelungsgeschichte des Pollens bei den Phanerogamen. Ovell & Füssli, Zürich.
  2. Harris H. 1999. The birth of the cell. Yale University Press. p138
  3. Hofmeister W. 1848. Bot. Zeit. 6, cols 425, 649, 670.
  4. Hofmeister W. 1849. Die Entstehung des Embryos der Phanerogamen. Friedrich Hofmeister, Leipzig.
  5. Mayr E. 1982. The growth of biological thought. Harvard. p677
  6. Flemming W. Zellsubstanz Kern und Zelltheilung. Vogel, Leipzig.
  7. Van Beneden E. 1883. Arch. Biol. 4.
  8. Harris H. 1999. The birth of the cell. Yale University Press. p141, 153
  9. Rabl C. 1885. Morphol. Jahrb. 10, 24.
  10. Coleman W. 1965. Cell, nucleus, and inheritance: an historical discovery. Proc. Am. Philos. Soc. 109, 124-158.
  11. Haeckel E. 1866. Generelle Morphologie der Organismen: Allgemeine Gründzuge der organischen Formen-Wissenschaft. 2 vols, Reimer, Berlin.
  12. Mayr E. 1982. The growth of biological thought. Harvard. p749
  13. Thanbichler M, Shapiro L (2006). Chromosome organization and segregation in bacteria. J. Struct. Biol. 156 (2): 292–303.
  14. Nakabachi A, Yamashita A, Toh H, Ishikawa H, Dunbar H, Moran N, Hattori M (2006). The 160-kilobase genome of the bacterial endosymbiont Carsonella. Science 314 (5797): 267.
  15. Pradella S, Hans A, Spröer C, Reichenbach H, Gerth K, Beyer S (2002). Characterisation, genome size and genetic manipulation of the myxobacterium Sorangium cellulosum So ce56. Arch Microbiol 178 (6): 484-92.
  16. Hinnebusch J, Tilly K (1993). Linear plasmids and chromosomes in bacteria. Mol Microbiol 10 (5): 917-22. PMID 7934868.
  17. Kelman LM, Kelman Z (2004). Multiple origins of replication in archaea. Trends Microbiol. 12 (9): 399–401.
  18. Thanbichler M, Wang SC, Shapiro L (2005). The bacterial nucleoid: a highly organized and dynamic structure. J. Cell. Biochem. 96 (3): 506–21.
  19. Sandman K, Pereira SL, Reeve JN (1998). Diversity of prokaryotic chromosomal proteins and the origin of the nucleosome. Cell. Mol. Life Sci. 54 (12): 1350–64.
  20. Sandman K, Reeve JN (2000). Structure and functional relationships of archaeal and eukaryal histones and nucleosomes. Arch. Microbiol. 173 (3): 165–9.
  21. Pereira SL, Grayling RA, Lurz R, Reeve JN (1997). Archaeal nucleosomes. Proc. Natl. Acad. Sci. U.S.A. 94 (23): 12633–7.
  22. 22.0 22.1 22.2 Dubcovsky J, Luo MC, Zhong GY, et al (1996). Genetic map of diploid wheat, Triticum monococcum L., and its comparison with maps of Hordeum vulgare L. Genetics 143 (2): 983–99.
  23. Bogin, Barry, Edward Alcamo, Curtis Chubb, William J. Ehmann, Mark R. Feil, David R. Hershey, Mitchell Leslie, Karel F. Liem, William Thwaites, and Salvatore Tocci. Austin: Holt, Rinehart, and Winston, 1999. 146.
  24. Umeko Semba, Yasuko Umeda, Yoko Shibuya, Hiroaki Okabe, Sumio Tanase and Tetsuro Yamamoto (2004). Primary structures of guinea pig high- and low-molecular-weight kininogens. International Immunopharmacology 4 (10-11): 1391-1400.
  25. Bogin, Barry, Edward Alcamo, Curtis Chubb, William J. Ehmann, Mark R. Feil, David R. Hershey, Mitchell Leslie, Karel F. Liem, William Thwaites, and Salvatore Tocci. Austin: Holt, Rinehart, and Winston, 1999. 146.
  26. 26.0 26.1 De Grouchy J (1987). Chromosome phylogenies of man, great apes, and Old World monkeys. Genetica 73 (1-2): 37–52.
  27. Houck ML, Kumamoto AT, Gallagher DS, Benirschke K (2001). Comparative cytogenetics of the African elephant (Loxodonta africana) and Asiatic elephant (Elephas maximus). Cytogenet. Cell Genet. 93 (3-4): 249–52.
  28. Wayne RK, Ostrander EA (1999). Origin, genetic diversity, and genome structure of the domestic dog. Bioessays 21 (3): 247–57.
  29. Burt DW (2002). Origin and evolution of avian microchromosomes. Cytogenet. Genome Res. 96 (1-4): 97–112.
  30. Ciudad J, Cid E, Velasco A, Lara JM, Aijón J, Orfao A (2002). Flow cytometry measurement of the DNA contents of G0/G1 diploid cells from three different teleost fish species. Cytometry 48 (1): 20–5.
  31. Yasukochi Y, Ashakumary LA, Baba K, Yoshido A, Sahara K (2006). A second-generation integrated map of the silkworm reveals synteny and conserved gene order between lepidopteran insects. Genetics 173 (3): 1319–28.
  32. Sakamura, T. (1918), Kurze Mitteilung uber die Chromosomenzahlen und die Verwandtschaftsverhaltnisse der Triticum-Arten. Bot. Mag., 32: 151-154.
  33. Charlebois R.L. (ed) 1999. Organization of the prokaryote genome. ASM Press, Washington DC.
  34. White M.J.D. 1973. The chromosomes. 6th ed, Chapman & Hall, London. p28
  35. von Winiwarter H. 1912. Études sur la spermatogenese humaine. Arch. biologie 27, 93, 147-9.
  36. Painter T.S. 1922. The spermatogenesis of man. Anat. Res. 23, 129.
  37. Painter T.S. 1923. Studies in mammalian spermatogenesis II. The spermatogenesis of man. J. Exp. Zoology 37, 291-336.
  38. Tjio J.H & Levan A. 1956. The chromosome number of man. Hereditas 42, 1-6.
  39. Hsu T.C. Human and mammalian cytogenetics: a historical perspective. Springer-Verlag, N.Y.
  40. http://vega.sanger.ac.uk/Homo_sapiens/index.html All data in this table was derived from this database, July 7 2007.
  41. Sequenced percentages are based on fraction of euchromatin portion, as the Human Genome Project goals called for determination of only the euchromatic portion of the genome. Telomeres, centromeres, and other heterochromatic regions have been left undetermined, as have a small number of unclonable gaps. See http://www.ncbi.nlm.nih.gov/genome/seq/ for more information on the Human Genome Project.


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