Karyotype - Knowledge

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fluoresces. Since regions rich in AT are more common in heterochromatin than in euchromatin, these regions are labelled preferentially. The different intensities of the single bands mirror the different contents of AT. Other fluorochromes like DAPI or Hoechst 33258 lead also to characteristic, reproducible patterns. Each of them produces its specific pattern. In other words: the properties of the bonds and the specificity of the fluorochromes are not exclusively based on their affinity to regions rich in AT. Rather, the distribution of AT and the association of AT with other molecules like histones, for example, influences the binding properties of the fluorochromes.

504: 1612:. The name is derived from centromeric or constitutive heterochromatin. The preparations undergo alkaline denaturation prior to staining leading to an almost complete depurination of the DNA. After washing the probe the remaining DNA is renatured again and stained with Giemsa solution consisting of methylene azure, methylene violet, methylene blue, and eosin. Heterochromatin binds a lot of the dye, while the rest of the chromosomes absorb only little of it. The C-bonding proved to be especially well-suited for the characterization of plant chromosomes. 1652: 57: 496: 72: 816: 1781: 1737: 4753: 874:(chromosome arm) is connected at the centromere, for a total of 4c. The chromosomes on micrographic karyograms are in this state as well, because they are generally micrographed in metaphase, but during this phase the two copies of each chromosome are so close to each other that they appear as one unless the image resolution is high enough to distinguish them. In reality, during the G 1803:. Using this technique, it is possible to detect small alterations in the human genome, that cannot be detected through methods employing metaphase chromosomes. Some loci deletions are known to be related to the development of cancer. Such deletions are found through digital karyotyping using the loci associated with cancer development. 247: 1764:. Because there are a limited number of spectrally distinct fluorophores, a combinatorial labeling method is used to generate many different colors. Fluorophore combinations are captured and analyzed by a fluorescence microscope using up to 7 narrow-banded fluorescence filters or, in the case of spectral karyotyping, by using an 936:

In our view, it is unlikely that one process or the other can independently account for the wide range of karyotype structures that are observed ... But, used in conjunction with other phylogenetic data, karyotypic fissioning may help to explain dramatic differences in diploid numbers between closely

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Differences in basic number of chromosomes. These differences could have resulted from successive unequal translocations which removed all the essential genetic material from a chromosome, permitting its loss without penalty to the organism (the dislocation hypothesis) or through fusion. Humans

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In many instances, endopolyploid nuclei contain tens of thousands of chromosomes (which cannot be exactly counted). The cells do not always contain exact multiples (powers of two), which is why the simple definition 'an increase in the number of chromosome sets caused by replication without cell

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for staining. The pattern of bands is very similar to that seen in G-banding. They can be recognized by a yellow fluorescence of differing intensity. Most part of the stained DNA is heterochromatin. Quinacrin (atebrin) binds both regions rich in AT and in GC, but only the AT-quinacrin-complex

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Chromosomes display a banded pattern when treated with some stains. Bands are alternating light and dark stripes that appear along the lengths of chromosomes. Unique banding patterns are used to identify chromosomes and to diagnose chromosomal aberrations, including chromosome breakage, loss,

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The results are clear. The inversions, when plotted in tree form (and independent of all other information), show a clear "flow" of species from older to newer islands. There are also cases of colonization back to older islands, and skipping of islands, but these are much less frequent. Using

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The polytene banding of the 'picture wing' group, the best-studied group of Hawaiian drosophilids, enabled Carson to work out the evolutionary tree long before genome analysis was practicable. In a sense, gene arrangements are visible in the banding patterns of each chromosome. Chromosome

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of each chromosome is generally shown for brevity, and in reality they are generally so close together that they look as one on photomicrographs as well unless the resolution is high enough to distinguish them. The study of whole sets of chromosomes is sometimes known as

1594:. It yields a series of lightly and darkly stained bands — the dark regions tend to be heterochromatic, late-replicating and AT rich. The light regions tend to be euchromatic, early-replicating and GC rich. This method will normally produce 300–400 bands in a normal, 1914:, believed to be the 4th most common trisomy, has many long lived affected individuals but only in a form other than a full trisomy, such as trisomy 9p syndrome or mosaic trisomy 9. They often function quite well, but tend to have trouble with speech. 1772:

in a dedicated image analysis software. Thus, chromosomes or chromosome sections can be visualized and identified, allowing for the analysis of chromosomal rearrangements. In the case of spectral karyotyping, image processing software assigns a

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have one pair fewer chromosomes than the great apes. Human chromosome 2 appears to have resulted from the fusion of two ancestral chromosomes, and many of the genes of those two original chromosomes have been translocated to other chromosomes.

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Although much is known about karyotypes at the descriptive level, and it is clear that changes in karyotype organization has had effects on the evolutionary course of many species, it is quite unclear what the general significance might be.

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The next stage took place after the development of genetics in the early 20th century, when it was appreciated that chromosomes (that can be observed by karyotype) were the carrier of genes. The term karyotype as defined by the

1217:) is also common, and some species of ferns have reached levels of polyploidy far in excess of the highest levels known in flowering plants. Polyploidy in animals is much less common, but it has been significant in some groups. 1066:

They simply could not believe what they saw ... They kept quiet for two or three years because they thought something was wrong with their tissue culture ... But when they obtained a couple more specimens they confirmed

1362:, where every number from x = 3 to x = 15 is represented by at least one species. Evidence of various kinds shows that trends of evolution have gone in different directions in different groups. In primates, the 994:, portions of the chromosomes are cast away in particular cells. This process is a carefully organised genome rearrangement where new telomeres are constructed and certain heterochromatin regions are lost. In 1601:

R-banding is the reverse of G-banding (the R stands for "reverse"). The dark regions are euchromatic (guanine-cytosine rich regions) and the bright regions are heterochromatic (thymine-adenine rich regions).

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it is always the paternal X which is inactivated. In human females some 15% of somatic cells escape inactivation, and the number of genes affected on the inactivated X chromosome varies between cells: in

921:, the same cannot be said for their karyotypes, which are highly variable. There is variation between species in chromosome number, and in detailed organization, despite their construction from the same 90:

is a graphical depiction of a karyotype, wherein chromosomes are generally organized in pairs, ordered by size and position of centromere for chromosomes of the same size. Karyotyping generally combines

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This process (especially studied in insects and some higher plants such as maize) may be a developmental strategy for increasing the productivity of tissues which are highly active in biosynthesis.

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is a technique used to quantify the DNA copy number on a genomic scale. Short sequences of DNA from specific loci all over the genome are isolated and enumerated. This method is also known as

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duplication, translocation or inverted segments. A range of different chromosome treatments produce a range of banding patterns: G-bands, R-bands, C-bands, Q-bands, T-bands and NOR-bands.

1135:, of a karyotype is the number of visible major chromosomal arms per set of chromosomes. Thus, FN ≤ 2 x 2n, the difference depending on the number of chromosomes considered single-armed ( 399:). In metaphase where the karyotype is typically studied, all DNA is condensed, but most of the time, DNA with a high GC content is usually less condensed, that is, it tends to appear as 1382:

for different chromosome structural forms. The structural variation may be associated with different numbers of chromosomes in different individuals, which occurs in the ladybird beetle

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The schematic karyogram in this section is a graphical representation of the idealized karyotype. For each chromosome pair, the scale to the left shows the length in terms of million

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Multicolor FISH is used to identify structural chromosome aberrations in cancer cells and other disease conditions when Giemsa banding or other techniques are not accurate enough.

1205:. The proportion of flowering plants which are polyploid was estimated by Stebbins to be 30–35%, but in grasses the average is much higher, about 70%. Polyploidy in lower plants ( 2080:

in 1922 was not certain whether the diploid of humans was 46 or 48, at first favoring 46, but revised his opinion from 46 to 48, and he correctly insisted on humans having an

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Differences in absolute sizes of chromosomes. Chromosomes can vary in absolute size by as much as twenty-fold between genera of the same family. For example, the legumes

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We have a very poor understanding of the causes of karyotype evolution, despite many careful investigations ... the general significance of karyotype evolution is obscure.

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Endopolyploidy and polyteny in differentiation and evolution: towards an understanding of quantitative and qualitative variation of nuclear DNA in ontogeny and phylogeny

2557: 1201:, where there are more than two sets of homologous chromosomes in the cells, occurs mainly in plants. It has been of major significance in plant evolution according to 4095:

Craddock E.M. (2000). "Speciation Processes in the Adaptive Radiation of Hawaiian Plants and Animals". In Hecht, Max K.; MacIntyre, Ross J.; Clegg, Michael T. (eds.).

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is the process by which a karyotype is discerned by determining the chromosome complement of an individual, including the number of chromosomes and any abnormalities.

1849:, in which three copies of a chromosome are present instead of the usual two, are common numerical abnormalities. Structural abnormalities often arise from errors in 5319: 598: 3396: 1701:-rich regions. Each chromosome has a characteristic banding pattern that helps to identify them; both chromosomes in a pair will have the same banding pattern. 586:, whereas the schematic karyogram shows the purple hue as typically seen on Giemsa stain (and is a result of its azure B component, which stains DNA purple). 250:

Micrograph of human chromosomes before further processing. Staining with Giemsa confers a purple color to chromosomes, but micrographs are often converted to

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attached to a fluorescence microscope. In the case of an mFISH image, every combination of fluorochromes from the resulting original images is replaced by a

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chromosome pairs (chromosomes 1 to 22). Homologous means that they have the same genes in the same loci, and autosomal means that they are not sex chromomes.

850:, making a total copy number of 2 for each locus (2c). At top center in the schematic karyogram, it also shows the chromosome 3 pair after having undergone 1724:

syndrome involves a deletion on the short arm of chromosome 5. It is written as 46,XX,5p-. The critical region for this syndrome is deletion of p15.2 (the

1928:(cry of the cat), from a truncated short arm on chromosome 5. The name comes from the babies' distinctive cry, caused by abnormal formation of the larynx. 523:

Both the micrographic and schematic karyograms shown in this section have a standard chromosome layout, and display darker and lighter regions as seen on

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species in Hawaiʻi have apparently descended from a single ancestral species that colonized the islands, probably 20 million years ago. The subsequent

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A full account of a karyotype may therefore include the number, type, shape and banding of the chromosomes, as well as other cytogenetic information.

238:. Karyograms are generally made by chromosomes in prometaphase or metaphase. During these phases, the two copies of each chromosome (connected at the 1853:. Both types of abnormalities can occur in gametes and therefore will be present in all cells of an affected person's body, or they can occur during 2762: 2707: 2654: 2542: 1704:

Karyotypes are arranged with the short arm of the chromosome on top, and the long arm on the bottom. Some karyotypes call the short and long arms

1043: 838:) which is the most common state of cells. The schematic karyogram in this section also shows this state. In this state (as well as during the G 4820: 1995: 1497: 4763: 2934:

Wyngaard GA, Gregory TR (December 2001). "Temporal control of DNA replication and the adaptive value of chromatin diminution in copepods".

1950:– 50% of cases have a segment of the long arm of chromosome 15 missing; a deletion of the paternal genes, example of imprinting disorder. 4781: 1554:

There are other animals and plants on the Hawaiian archipelago which have undergone similar, if less spectacular, adaptive radiations.

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such as an extra chromosome or one or more chromosomes lost. Abnormalities in chromosome number usually cause a defect in development.

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Searle, J. B. (1 June 1984). "Three New Karyotypic Races of the Common Shrew Sorex Araneus (Mammalia: Insectivora) and a Phylogeny".

806:), although this is not included in micrographic karyograms in clinical practice. Its genome is relatively tiny compared to the rest. 1077:

The number of chromosomes in the karyotype between (relatively) unrelated species is hugely variable. The low record is held by the

870:(annotated as "Meta."). During this interval, there is still 2n, but each chromosome will have 2 copies of each locus, wherein each 5217: 1642:-associated protein. This yields a dark region where the silver is deposited, denoting the activity of rRNA genes within the NOR. 1335:

is the condition in which the chromosome number in the cells is not the typical number for the species. This would give rise to a

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Alternatively, the human genome can be classified as follows, based on pairing, sex differences, as well as location within the

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Gustavsson, Ingemar (3 March 1969). "Cytogenetics, distribution and phenotypic effects of a translocation in Swedish cattle".

1016:, the inactivation is random as between the two Xs; thus the mammalian female is a mosaic in respect of her X chromosomes. In 5309: 4950: 4197: 4164: 4139: 4112: 3596: 2053:

Investigation into the human karyotype took many years to settle the most basic question: how many chromosomes does a normal

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Differences in relative size of chromosomes. These differences probably arose from segmental interchange of unequal lengths.

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Pellicciari, C.; Formenti, D.; Redi, C.A.; Manfredi, M.G.; Romanini (February 1982). "DNA content variability in primates".

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Carrel L, Willard H (2005). "X-inactivation profile reveals extensive variability in X-linked gene expression in females".

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The work took place in 1955, and was published in 1956. The karyotype of humans includes only 46 chromosomes. The other

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Liehr T, Starke H, Weise A, Lehrer H, Claussen U (January 2004). "Multicolour FISH probe sets and their applications".

4056: 1745: 17: 2507:"Relevance of human chromosome analysis activities against mutation concept in genetics course. IOP Conference Series" 4813: 3827: 3423: 2489: 2380: 2290: 2223: 1940:– 50% of cases have a segment of the long arm of chromosome 15 missing; a deletion of the maternal genes, example of 1664: 1457: 4240:

Schröck E, du Manoir S, Veldman T, et al. (July 1996). "Multicolor spectral karyotyping of human chromosomes".

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Chromosome abnormalities can be numerical, as in the presence of extra or missing chromosomes, or structural, as in

1712:, respectively. In addition, the differently stained regions and sub-regions are given numerical designations from 1437:

can reveal relationships between closely related species: the classic example is the study of chromosome banding in

771:(in green rectangle at bottom right in the schematic karyogram, with adjacent silhouettes of typical representative 5089: 4769: 3205:

Kim, D.S.; Nam, Y.K.; Noh, J.K.; Park, C.H.; Chapman, F.A. (2005). "Karyotype of North American shortnose sturgeon

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MJ, Kottler (1974). "From 48 to 46: cytological technique, preconception, and the counting of human chromosomes".

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Judd, Walter S.; Campbell, Christopher S.; Kellogg, Elizabeth A.; Stevens, Peter F.; Donoghue, Michael J. (2002).

846:), each cell has 2 autosomal chromosomes of each kind (designated 2n), where each chromosome has one copy of each 4775: 4663: 1521: 115:

karyogram is a designed graphic representation of a karyotype. In schematic karyograms, just one of the sister

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to each spectrally different combination, allowing the visualization of the individually colored chromosomes.

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Aneuploidy may also occur within a group of closely related species. Classic examples in plants are the genus

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Polyploid series in related species which consist entirely of multiples of a single basic number are known as

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In some cases there is even significant variation within species. In a review, Godfrey and Masters conclude:

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chromosomes are many times larger. These differences probably reflect different amounts of DNA duplication.

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are another example, though in this case they would not be regarded as normal members of the population.

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Bangs, 1900 (Rodentia, Sigmodontinae) of the Brazilian Cerrado, with the description of two new species"

3289:"Karyotype characterization and ZZ/ZW sex chromosome heteromorphism in two species of the catfish genus 4074:"Chromosomes and male genitalia of Hawaiian Drosophila: tools for interpreting phylogeny and geography" 3328: 2192: 1309: 1979: 5082: 3991: 1822: 2187: 5136: 4920: 2323:

Mouse Genetics, Concepts and Applications. Chapter 5.2: KARYOTYPES, CHROMOSOMES, AND TRANSLOCATIONS

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Comprehensive Sampling and Sample Preparatio. Chapter: 3.05 - Blood Sample Collection and Handling

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Fusion of ancestral chromosomes left distinctive remnants of telomeres, and a vestigial centromere

3297: 1958: 540: 1004:. The inactivation of one X chromosome takes place during the early development of mammals (see 5042: 5015: 4560:. Cambridge University Press. 2nd ed, revised and enlarged, 1958. Oliver & Boyd, Edinburgh. 2372: 2248: 1931: 1760:

labeled probes for each chromosome are made by labeling chromosome-specific DNA with different

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stain to adhere correctly, all chromosomal proteins must be digested and removed. For humans,

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Also documented are trisomy 8 and trisomy 16, although they generally do not survive to birth.

5352: 5342: 5077: 4975: 4081: 2756: 2701: 2648: 2536: 1947: 1882:, the most common male chromosomal disease, otherwise known as 47,XXY, is caused by an extra 1826: 1818: 1540: 1481: 1160: 1156: 1152: 1148: 1144: 758: 622: 476: 143:, and any other physical characteristics. The preparation and study of karyotypes is part of 2353:

2nd ed, ed. M.J. Barch. The Association of Cytogenetic Technologists, Raven Press, New York.

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female to colonise an island, it is more likely to have been a group from the same species.

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to facilitate data presentation and make comparisons of results from different laboratories.

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Zelenin, A. V.; Rodionov, A. V.; Bolsheva, N. L.; Badaeva, E. D.; Muravenko, O. V. (2016).

4304: 4249: 3956: 3768: 3218: 3136: 3065: 3010: 2943: 2789: 2602: 2321: 2157: 1879: 1082: 736: 694: 503: 373: 1143:) present. Humans have FN = 82, due to the presence of five acrocentric chromosome pairs: 961:

Instead of the usual gene repression, some organisms go in for large-scale elimination of

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Matthey, R. (15 May 1945). "L'evolution de la formule chromosomiale chez les vertébrés".

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cells of an otherwise genetically normal individual; one well-documented example is the

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is also acrocentric). The fundamental autosomal number or autosomal fundamental number,

4701: 4684: 4499: 4442: 4387: 4273: 4020: 3856: 3711: 3519: 3465: 3269: 3234: 3191: 3160: 3089: 3034: 2916: 2873: 2739: 2722: 2631: 2604: 2585: 2524: 2427: 2400: 2365: 2279: 2145: 2102:'s lab found the chromosome count to be 46 using new techniques available at the time: 2077: 1941: 1536: 1509: 1489: 1367: 1107: 532: 507:

Schematic karyogram of a human. Even at low magnification, it gives an overview of the

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Leary, Rebecca J; Cummins, Jordan; Wang, Tian-Li; Velculescu, Victor E (August 2007).

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is now known to be a result of an end-to-end fusion of two ancestral ape chromosomes.

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de Oliveira, R.R.; Feldberg, E.; dos Anjos, M. B.; Zuanon, J. (July–September 2007).

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Cutting up a photomicrograph and arranging the result into an indisputable karyogram.

2023: 1937: 1800: 1651: 1460:. These roughly 800 Hawaiian drosophilid species are usually assigned to two genera, 1384: 1279: 1092: 1058: 1048: 1013: 483: 458: 300: 242:) will look as one unless the image resolution is high enough to distinguish the two. 151: 4503: 4391: 4277: 3715: 3469: 3380: 3311: 3288: 3273: 3238: 3164: 3093: 2920: 2891:

Müller F, Bernard V, Tobler H (February 1996). "Chromatin diminution in nematodes".

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means that chromosome number can vary even within one interbreeding population; and

5116: 4995: 4766:, an online activity from the University of Utah's Genetic Science Learning Center. 4696: 4644: 4543: 4539: 4530:

Battaglia, Emilio (1994). "Nucleosome and nucleotype: a terminological criticism".

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Souza, A.L.G.; de O. Corrêa, M.M.; de Aguilar, C.T.; Pessôa, L.M. (February 2011).

3354: 3337: 3306: 3261: 3226: 3187: 3144: 3073: 3038: 3018: 2951: 2900: 2857: 2807: 2797: 2734: 2681: 2626: 2616: 2569: 2514: 2422: 2412: 2047: 2043: 2031: 1987: 1889: 1725: 1449: 1356:, where the gametic (= haploid) numbers form the series x = 3, 4, 5, 6, and 7; and 1290: 1140: 1136: 871: 847: 594: 92: 3968: 3148: 3077: 2133:

Squashing the preparation on the slide forcing the chromosomes into a single plane

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or in an individual organism, mainly including their sizes, numbers, and shapes.

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Comai L (November 2005). "The advantages and disadvantages of being polyploid".

2113: 1829:, large-scale deletions or duplications. Numerical abnormalities, also known as 1452:

have the most diverse collection of drosophilid flies in the world, living from

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Adams KL, Wendel JF (April 2005). "Polyploidy and genome evolution in plants".

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system. Considering the techniques of the time, these results were remarkable.

1905: 1858: 1834: 1765: 1635: 1409: 1247: 1001: 977: 768: 578:(sperm and egg cells) which are haploid with 23 unpaired chromosomes, and this 548: 359: 331:

Six different characteristics of karyotypes are usually observed and compared:

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are used most frequently because they are easily induced to divide and grow in

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ahead with an average of 1262 chromosomes. Top score for animals might be the

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Schematic karyograms generally display a DNA copy number corresponding to the

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Comparative karyological study of species Muscari Mill. and Bellevalia Lapeyr

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Goday C, Esteban MR (March 2001). "Chromosome elimination in sciarid flies".

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A preparation which includes the dyes Methylene Blue, Eosin Y and Azure-A,B,C

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is not shown in these karyograms. The micrographic karyogram is converted to

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Stebbins, G.L. (1940). "The significance of polyploidy in plant evolution".

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Some disorders arise from loss of just a piece of one chromosome, including

1175:, of a karyotype is the number of visible major chromosomal arms per set of 1034:

A spectacular example of variability between closely related species is the

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and does not show visually distinguishable chromosomes even on micrography.

4955: 4829: 4668: 4648: 4495: 4438: 4383: 4367: 4336: 4317: 4226: 4029: 3707: 3666: 3657: 3636: 3558: 3515: 3156: 3030: 2963: 2904: 2869: 2821: 2748: 2640: 2581: 2436: 2099: 2095: 1660: 1595: 1587: 1576: 1544: 1263: 1259: 1164: 1102: 1056:. When they looked at the karyotype of the closely related Indian muntjac, 990: 926: 925:. This variation provides the basis for a range of studies in evolutionary 788: 780: 750: 746: 536: 508: 337: 292: 246: 213: 201: 144: 4583: 4269: 4071: 3976: 3864: 3800: 3178:

Khandelwal S. (1990). "Chromosome evolution in the genus Ophioglossum L".

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employs several techniques to visualize different aspects of chromosomes:

71: 5174: 5169: 5032: 5010: 5005: 4873: 4710: 2504: 1925: 1774: 1769: 1761: 1757: 1749: 1736: 1721: 1616: 1275: 1267: 1241: 1053: 400: 355: 218: 96: 42: 3022: 2778:"Kinetochore reproduction theory may explain rapid chromosome evolution" 2042:. The subsequent history of the concept can be followed in the works of 1685:, is used to stain bands on the chromosomes. Giemsa is specific for the 1504:, which can be dated to 30 mya. The archipelago itself (produced by the 1500:). The oldest member of the Hawaiian archipelago still above the sea is 1062:, they were astonished to find it had female = 6, male = 7 chromosomes. 5194: 5184: 5179: 5164: 5104: 5094: 4970: 4944: 4895: 4885: 4833: 3361:(Rodentia: Sigmodontinae) from Chapada Diamantina, northeastern Brazil" 3265: 2141: 2128: 2019: 1983: 1830: 1753: 1656: 1620: 1609: 1513: 1501: 1468: 1462: 1422: 1363: 1332: 1214: 1210: 1198: 1022: 1017: 985: 918: 843: 835: 820: 815: 618: 606: 590: 566:

within a normal cell of the human body, and which contains 22 pairs of

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from Biology Corner, a resource site for biology and science teachers.

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Carson HL (June 1970). "Chromosome tracers of the origin of species".

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10.1002/1521-1878(200103)23:3<242::AID-BIES1034>3.0.CO;2-P

2218:(2 ed.). Sunderland MA, US: Sinauer Associates Inc. p. 544. 1902:, a common chromosomal disease, is caused by trisomy of chromosome 21. 1370:

was formed by a merger of ancestral chromosomes, reducing the number.

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karyogram demonstrating the basic knowledge needed to read a karyotype

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ISCN 2005: An International System for Human Cytogenetic Nomenclature

3757:"Origin of human chromosome 2: an ancestral telomere-telomere fusion" 2955: 2720: 2667: 2401:"Analytical Biases Associated with GC-Content in Molecular Evolution" 2124: 1999: 1911: 1686: 1682: 1583: 1493: 1414: 1301: 1120: 1005: 970: 922: 883: 867: 803: 772: 761: 583: 558:

Both the micrographic and schematic karyograms show the normal human

539:. Compared to darker regions, the lighter regions are generally more 524: 499:

Micrographic karyogram of a human male. See section text for details.

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Gomes CJ, Harman MW, Centuori SM, Wolgemuth CW, Martinez JD (2018).

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dating, the present islands date from 0.4 million years ago (mya) (

1418: 1305: 1180: 1176: 1078: 981: 894: 859: 827: 792: 791:

denoted 46,XY. However, approximately 0.018% percent of humans are

629:), the human chromosomes are classified into the following groups: 567: 424: 388: 264: 209: 31: 4597:

von Winiwarter H. (1912). "Études sur la spermatogenèse humaine".

3813: 3755:

IJdo JW, Baldini A, Ward DC, Reeders ST, Wells RA (October 1991).

527:, which is the appearance of the chromosomes after treatment with 5302: 5156: 5067: 5062: 4786: 4291:

Wang TL, Maierhofer C, Speicher MR, et al. (December 2002).

3992:"Chromosomal sequences and interisland colonizations in Hawaiian 3927:"The Hawaiian-Emperor volcanic chain, Part I. Geologic evolution" 2120: 2066: 2054: 1991: 1893: 1869:

Chromosomal abnormalities that lead to disease in humans include

1854: 1846: 1838: 1698: 1694: 1591: 1405: 1389: 1255: 1237: 1233: 1222: 1087: 1035: 1025:

cells up about 25% of genes on the Barr body escape inactivation.

855: 776: 559: 528: 462: 420: 396: 392: 384: 235: 181:

chromosomes are present in two copies. There may, or may not, be

174: 46: 4635:

Painter T.S. (1923). "Studies in mammalian spermatogenesis II".

3818:

A glossary of genetics and cytogenetics: Classical and molecular

2605:"Measuring DNA content in live cells by fluorescence microscopy" 1286:, the daughter chromosomes separating from each other inside an 617:

Based on the karyogram characteristics of size, position of the

5212: 5099: 4863: 4859: 4854: 4752: 3204: 1954: 1842: 1675: 1548: 1394: 1358: 1352: 1191: 998:, all the somatic cell precursors undergo chromatin diminution. 858:(annotated as S) of the cell cycle. This interval includes the 579: 575: 563: 432: 296: 272: 190: 64: 4404: 3293:

Kner, 1854 (Siluriformes: Loricariidae) from the Amazon basin"

1732:

Multicolor FISH (mFISH) and spectral karyotype (SKY technique)

511:, with numbered chromosome pairs, its main changes during the 5277: 5262: 5257: 5252: 5247: 5242: 5237: 5232: 5227: 5202: 5140: 4187: 3209:

with the highest chromosome number in the Acipenseriformes".

1752:

techniques used to simultaneously visualize all the pairs of

1728:

on the chromosome), which is written as 46,XX,del(5)(p15.2).

1517: 1484:, make it possible to see which species are closely related. 893:

per human cell varies from 0 (erythrocytes) up to 1,500,000 (

784: 519:

to scale (at bottom left). See section text for more details.

312: 27:

Photographic display of total chromosome complement in a cell

4724:

Human chromosome 2 is a fusion of two ancestral. chromosomes

3122: 1994:, in 1882. The name was coined by another German anatomist, 897:), mainly depending on the number of mitochondria per cell. 3924: 2027: 1206: 1097: 593:, and the scale to the right shows the designations of the 376:

are small bodies attached to a chromosome by a thin thread.

196:

Karyotypes can be used for many purposes; such as to study

4349: 4078:

Hawaiian biogeography: evolution on a hot spot archipelago

3111:

Human and mammalian cytogenetics: a historical perspective

2213: 4080:. Washington DC: Smithsonian Institution Press. pp. 1934:, from the loss of part of the short arm of chromosome 1. 1690: 1671: 1433:

The detailed study of chromosome banding in insects with

1194:

is the number of complete sets of chromosomes in a cell.

965:, or other kinds of visible adjustment to the karyotype. 914: 315:

can be predicted by observation of interphase cells (see

268: 135:. Attention is paid to their length, the position of the 3932:. In Decker, R.W.; Wright, T.L.; Stauffer, P.H. (eds.). 1861:

individual who has some normal and some abnormal cells.

1323:

for the investigation of ancient karyotype duplications.

900: 574:(allosomes). A major exception to diploidy in humans is 5320:

International System for Human Cytogenetic Nomenclature

4664:"Joe Hin Tjio The man who cracked the chromosome count" 4239: 4072:

Kaneshiro, K.Y.; Gillespie, R.G.; Carson, H.L. (1995).

3052:

Wurster DH, Benirschke K (June 1970). "Indian muntjac,

2168:

Pages displaying short descriptions of redirect targets

973:, entire chromosomes are eliminated during development. 599:

International System for Human Cytogenomic Nomenclature

4290: 4212: 3754: 3634: 3394: 2989: 2238: 2160: – Symbols and abbreviations used in cytogenetics 1115:

at 372 chromosomes. The existence of supernumerary or

307:. Sometimes observations may be made on non-dividing ( 2723:"Mitochondria in oocyte aging: current understanding" 2277:

Stebbins, G.L. (1950). "Chapter XII: The Karyotype".

2239:

King, R.C.; Stansfield, W.D.; Mulligan, P.K. (2006).

1448:

In about 6,500 sq mi (17,000 km), the

3620:(6th ed.). London: Chapman & Hall. p. 2980:. Sinauer Associates, Stamford CT. 8th ed, Chapter 9 2345: 2343: 1961:, a translocation mutation commonly associated with 969:

Chromosome elimination. In some species, as in many

937:

related species, which were previously inexplicable.

4616:Painter T.S. (1922). "The spermatogenesis of man". 2890: 2452:

Thompson & Thompson Genetics in Medicine 7th Ed

2398: 1876:

results from a single X chromosome (45,X or 45,X0).

605:. Each row of chromosomes is vertically aligned at 562:karyotype, which is the typical composition of the 372:Differences in number and position of satellites. 131:and what these chromosomes look like under a light 111:(or simply micrographic) karyogram. In contrast, a 30:"Idiogram" redirects here. Not to be confused with 3815: 3613: 2670:"Mitochondrial DNA in anucleate human blood cells" 2505:Erwinsyah, R., Riandi, & Nurjhani, M. (2017). 2364: 2278: 1978:Chromosomes were first observed in plant cells by 1598:. It is the most common chromosome banding method. 1300:The phenomenon occurs sporadically throughout the 834:of the cellular state (outside of the replicative 154:cells of an individual or a species is called the 4772:from the University of Arizona's Biology Project. 4682: 4596: 3051: 2340: 2247:(7th ed.). Oxford University Press. p. 1748:and the older spectral karyotyping are molecular 1646: 1308:to humans; it is diverse and complex, and serves 1096:. The high record would be somewhere amongst the 1029: 795:, sometimes due to variations in sex chromosomes. 358:. These differences probably came about through 41:is the general appearance of the complete set of 5334: 4131:Hawaiian natural history, ecology, and evolution 3056:: a deer with a low diploid chromosome number". 2394: 2392: 1512:) has existed for far longer, at least into the 1366:have 24x2 chromosomes whereas humans have 23x2. 419:. Euchromatin regions contain larger amounts of 4736:Evidence of common ancestry: human chromosome 2 3814:Rieger, R.; Michaelis, A.; Green, M.M. (1968). 3679: 3637:"New insights into cell cycle control from the 3395:Weksler, M.; Bonvicino, C.R. (3 January 2005). 2933: 2116:, which swells them and spreads the chromosomes 1973: 139:, banding pattern, any differences between the 4460:Vermeesch, Joris Robert; Rauch, Anita (2006). 4459: 4188:Lisa G. Shaffer; Niels Tommerup, eds. (2005). 4051:. Boca Raton FL: CRC Press. pp. 407–439. 2775: 2721:Zhang D, Keilty D, Zhang ZF, Chian RC (2017). 2668:Shuster RC, Rubenstein AJ, Wallace DC (1988). 1404:. There is some evidence from the case of the 189:cells have multiple copies of chromosomes and 4828: 4814: 4094: 3177: 3127:, an ant with only one pair of chromosomes". 3000: 2448: 2446: 2389: 2319: 1547:. Although it would be possible for a single 1282:) chromosomes in a 'resting' nucleus undergo 1046:. The diploid number of the Chinese muntjac, 956: 602: 439:and therefore produces the typical "G-Bands". 225: 4634: 4615: 4043:Carson H.L. (1992). "Inversions in Hawaiian 3682:"Endoreplication cell cycles: more for less" 3582: 2847: 2761:: CS1 maint: multiple names: authors list ( 2714: 2706:: CS1 maint: multiple names: authors list ( 2653:: CS1 maint: multiple names: authors list ( 2596: 2541:: CS1 maint: multiple names: authors list ( 2010:, "kernel", "seed", or "nucleus", and τύπος 1953:Chromosomal abnormalities can also occur in 263:The study of karyotypes is made possible by 4407:"Genome: Origins and evolution of the term" 4154: 4042: 3536: 3319: 2661: 2349:Gustashaw K.M. 1991. Chromosome stains. In 1806: 1373: 1254:tissues the cells have ceased to divide by 597:. Such bands and sub-bands are used by the 170:(humans: n = 23). Thus, in humans 2n = 46. 4821: 4807: 4782:Chromosome Staining and Banding Techniques 4047:". In Krimbas, C.B.; Powell, J.R. (eds.). 3842: 3408:Arquivos do Museu Nacional, Rio de Janeiro 2443: 2216:Plant systematics, a phylogenetic approach 1566: 1421:coast, that the two chromosome morphs are 612: 531:(to partially digest the chromosomes) and 379:Differences in degree and distribution of 4787:Bjorn Biosystems for Karyotyping and FISH 4700: 4529: 4477: 4326: 4316: 4155:Maloy, Stanley R.; Hughes, Kelly (2013). 4019: 3989: 3946: 3790: 3780: 3697: 3656: 3379: 3310: 2811: 2801: 2738: 2630: 2620: 2518: 2460: 2458: 2426: 2416: 1670:In the "classic" (depicted) karyotype, a 735:Very small, acrocentric (and 21, 22 with 166:(the sex cells) the chromosome number is 4770:Karyotyping activity with case histories 3571: 3482: 3447: 3280: 3180:Botanical Journal of the Linnean Society 3123:Crosland M.W.J.; Crozier, R.H. (1986). " 2362: 2276: 2260: 2258: 2086: 1779: 1735: 1650: 1590:following digestion of chromosomes with 1516:. Previous islands now beneath the sea ( 814: 502: 494: 347:each have six pairs of chromosomes, yet 245: 229: 70: 55: 4127: 3493: 3388: 3251: 3245: 3198: 976:Chromatin diminution (founding father: 150:The basic number of chromosomes in the 14: 5335: 4717: 4661: 3925:Clague, D.A.; Dalrymple, G.B. (1987). 3877: 3743:Chromosomal evolution in higher plants 3105: 3103: 2776:Godfrey LR, Masters JC (August 2000). 2477: 2455: 2367:Chromosomal evolution in higher plants 2351:The ACT Cytogenetics Laboratory Manual 1908:is caused by trisomy of chromosome 13. 1791: 1720:on the chromosome arms. For example, 295:when most condensed. In order for the 5310:List of organisms by chromosome count 4802: 4726:Alec MacAndrew; accessed 18 May 2006. 3912: 3628: 3611: 3348: 2468:. 3rd ed, Cambridge University Press. 2264: 2255: 1557: 1126: 901:Diversity and evolution of karyotypes 709:Small, metacentric or submetacentric 654:Large, metacentric or submetacentric 212:and to gather information about past 4076:. In Wagner, W.L.; Funk, E. (eds.). 3680:Edgar BA, Orr-Weaver TL (May 2001). 3591:. Academic Press. pp. 427–517. 2990:King, Stansfield & Mulligan 2006 1756:in an organism in different colors. 1740:Spectral karyogram of a human female 1240:. It is a common arrangement in the 882:phases, nuclear DNA is dispersed as 407:. GC rich DNA tends to contain more 4662:Wright, Pearce (11 December 2001). 3583:Gregory, T.R.; Mable, B.K. (2011). 3397:"Taxonomy of pygmy rice rats genus 3100: 2555: 2207: 1982:in 1842. Their behavior in animal ( 1571: 24: 4702:10.1111/j.1601-5223.1956.tb03010.x 4569: 4466:European Journal of Human Genetics 4157:Brenner's Encyclopedia of Genetics 3857:10.1111/j.1601-5223.1969.tb02259.x 3192:10.1111/j.1095-8339.1990.tb01876.x 2026:chromosomes, in contrast to their 980:). In this process, found in some 775:): The most common karyotypes for 490: 486:or otherwise abnormal individuals. 25: 5374: 4745: 4159:. San Diego, CA: Academic Press. 4049:Drosophila inversion polymorphism 3635:Lilly M.A.; Duronio R.J. (2005). 2511:Materials Science and Engineering 2328:Revised August 2004, January 2008 2281:Variation and evolution in plants 468:between members of a population ( 234:Chromosomes at various stages of 4751: 1896:(three copies) of chromosome 18. 1428: 1296:division' is not quite accurate. 693:Medium-sized, acrocentric, with 621:and sometimes the presence of a 543:active, with a greater ratio of 431:). The staining technique using 427:pairs (that is, it has a higher 4931:Macrochromosome/Microchromosome 4729: 4676: 4655: 4628: 4609: 4590: 4563: 4550: 4523: 4510: 4453: 4398: 4343: 4284: 4233: 4206: 4181: 4148: 4121: 4099:. Vol. 31. pp. 1–43. 4088: 4065: 4036: 3983: 3940: 3918: 3906: 3871: 3836: 3807: 3748: 3741:Stebbins, G. Ledley, Jr. 1972. 3735: 3722: 3673: 3605: 3576: 3565: 3530: 3487: 3476: 3441: 3381:10.1590/S1984-46702011000100013 3312:10.1590/S1679-62252007000300010 3171: 3116: 3045: 2994: 2983: 2970: 2927: 2884: 2841: 2828: 2769: 2549: 2498: 2471: 2356: 1262:contain more than the original 1100:, with the adder's tongue fern 354:Differences in the position of 326: 129:chromosome count of an organism 4685:"The chromosome number of man" 4544:10.1080/00087114.1994.10797297 4134:. University of Hawaii Press. 3585:"Ch. 8: Polyploidy in animals" 2520:10.1088/1757-899x/180/1/012285 2331: 2313: 2299: 2270: 2232: 2180: 2034:who worked with Lev Delaunay, 1655:Karyogram from a human female 1647:Classic karyotype cytogenetics 1030:Number of chromosomes in a set 810: 311:) cells. The sex of an unborn 63:karyogram of human male using 13: 1: 4462:"Reply to Hochstenbach et al" 4192:. Switzerland: S. Karger AG. 3969:10.1126/science.168.3938.1414 3822:. New York: Springer-Verlag. 3699:10.1016/S0092-8674(01)00334-8 3587:. In Gregory, T. Ryan (ed.). 3342:10.1016/S0047-2484(82)80045-6 3149:10.1126/science.231.4743.1278 3078:10.1126/science.168.3937.1364 2686:10.1016/s0006-291x(88)81291-9 2466:Animal cytology and evolution 2285:. Columbia University Press. 2173: 2057:human cell contain? In 1912, 1833:, often occur as a result of 1640:nucleolar organization region 1327: 787:usually have both an X and a 680:Medium-sized, submetacentric 4683:Tjio J.H.; Levan A. (1956). 4558:Evolution of genetic systems 4297:Proc. Natl. Acad. Sci. U.S.A 4262:10.1126/science.273.5274.494 3761:Proc. Natl. Acad. Sci. U.S.A 2782:Proc. Natl. Acad. Sci. U.S.A 2399:Romiguier J, Roux C (2017). 1974:History of karyotype studies 1967:acute lymphoblastic leukemia 1963:chronic myelogenous leukemia 1864: 1606:constitutive heterochromatin 1038:, which was investigated by 570:chromosomes and one pair of 415:. GC rich DNA is lighter on 7: 4776:Printable karyotype project 4105:10.1007/978-1-4615-4185-1_1 3589:The Evolution of the Genome 2371:. London: Arnold. pp. 2151: 2061:reported 47 chromosomes in 2030:contents was introduced by 1604:C-banding: Giemsa binds to 1480:rearrangements, especially 1244:, and in some other groups. 258: 10: 5379: 4971:Dinoflagellate chromosomes 4012:10.1093/genetics/103.3.465 3329:Journal of Human Evolution 2674:Biochem Biophys Res Commun 2326:. Oxford University Press. 2193:Collins English Dictionary 2166: – Laboratory process 1810: 1693:. Quinacrine binds to the 1052:, was found to be 46, all 957:Changes during development 917:is highly standardized in 891:human mitochondrial genome 800:human mitochondrial genome 465:and the rest of the body), 446:Variation is often found: 279:have been arrested during 226:Observations on karyotypes 193:cells have single copies. 29: 5315:List of sequenced genomes 5290: 5193: 5155: 5125: 5083:Chromosomal translocation 5053: 4956:A chromosome/B chromosome 4947:(or accessory chromosome) 4909: 4840: 4423:10.1134/S0026893316040178 4128:Ziegler, Alan C. (2002). 3551:10.1016/j.pbi.2005.01.001 3231:10.1007/s10228-004-0257-z 2622:10.1186/s13008-018-0039-z 2574:10.1080/00224490209552139 2558:"How Common is Intersex?" 1986:) cells was described by 1539:was spurred by a lack of 1186: 601:to describe locations of 477:geographic specialization 5137:Telomere-binding protein 4951:Supernumerary chromosome 3990:Carson HL (March 1983). 2838:. 2nd ed, Oxford. p218-9 2418:10.3389/fgene.2017.00016 2243:A dictionary of genetics 1851:homologous recombination 1813:Chromosome abnormalities 1807:Chromosome abnormalities 1784:Spectral human karyotype 1374:Chromosomal polymorphism 1131:The fundamental number, 722:Very small, metacentric 603:chromosome abnormalities 413:transcriptionally active 127:Karyotypes describe the 4479:10.1038/sj.ejhg.5201663 3892:10.1093/sysbio/33.2.184 3782:10.1073/pnas.88.20.9051 3450:The American Naturalist 3298:Neotropical Ichthyology 3211:Ichthyological Research 2834:Maynard Smith J. 1998. 2803:10.1073/pnas.97.18.9821 2562:Journal of Sex Research 2363:Stebbins, G.L. (1971). 2188:"Karyotype, definition" 2112:Pretreating cells in a 1980:Carl Wilhelm von Nägeli 1959:Philadelphia chromosome 1619:pattern obtained using 1567:Depiction of karyotypes 1425:to different habitats. 889:The copy number of the 783:and are denoted 46,XX; 625:(a segment distal to a 613:Human chromosome groups 470:chromosome polymorphism 198:chromosomal aberrations 5073:Structural alterations 4649:10.1002/jez.1400370303 4556:Darlington C.D. 1939. 4368:10.1038/nprot.2007.276 4318:10.1073/pnas.202610899 3658:10.1038/sj.onc.1208610 3612:White, M.J.D. (1973). 3539:Curr. Opin. Plant Biol 3207:Acipenser brevirostrum 3113:. Springer-Verlag, NY. 2905:10.1002/bies.950180209 2320:Lee M. Silver (1995). 2092: 1932:1p36 Deletion syndrome 1841:in the formation of a 1785: 1741: 1683:mepacrine (quinacrine) 1667: 1543:and a wide variety of 1522:Emperor Seamount Chain 1347:are examples of this. 1337:chromosome abnormality 1181:sex-linked chromosomes 1112:Acipenser brevirostrum 1075: 954: 939: 823: 667:Large, submetacentric 627:secondary constriction 520: 515:(top center), and the 500: 267:. Usually, a suitable 255: 243: 79: 68: 5090:Numerical alterations 5078:Chromosomal inversion 4976:Homologous chromosome 4352:"Digital karyotyping" 4293:"Digital karyotyping" 3745:. Nelson, London. p18 3732:. Elsevier, New York. 3359:Wiedomys pyrrhorhinus 2978:Developmental biology 2836:Evolutionary genetics 2727:Facts Views Vis Obgyn 2144:have 48 chromosomes. 2090: 1996:Heinrich von Waldeyer 1948:Prader-Willi syndrome 1819:derivative chromosome 1783: 1739: 1654: 1627:T-banding: visualize 1439:Hawaiian drosophilids 1398:, the European shrew 1250:occurs when in adult 1064: 944: 934: 818: 623:chromosomal satellite 506: 498: 249: 233: 74: 59: 5358:Evolutionary biology 5298:Extrachromosomal DNA 4986:Satellite chromosome 4961:Lampbrush chromosome 4901:Nuclear organization 4792:12 June 2019 at the 4760:at Wikimedia Commons 4599:Archives de Biologie 4097:Evolutionary Biology 3357:"A new karyotype of 2158:Cytogenetic notation 1990:, the discoverer of 1965:and less often with 1880:Klinefelter syndrome 1435:polytene chromosomes 1083:Parascaris univalens 517:mitochondrial genome 5363:Genetics techniques 4991:Centromere position 4966:Polytene chromosome 4936:Circular chromosome 4309:2002PNAS...9916156W 4254:1996Sci...273..494S 4215:Histol. Histopathol 3961:1970Sci...168.1414C 3934:Volcanism in Hawaii 3773:1991PNAS...88.9051I 3254:Experientia (Basel) 3223:2005IchtR..52...94K 3141:1986Sci...231.1278C 3070:1970Sci...168.1364W 3023:10.1038/nature03479 3015:2005Natur.434..400C 2976:Gilbert S.F. 2006. 2948:2001JEZ...291..310W 2794:2000PNAS...97.9821G 2464:White M.J.D. 1973. 2059:Hans von Winiwarter 1857:and give rise to a 1801:virtual karyotyping 1797:Digital karyotyping 1792:Digital karyotyping 1232:, where one sex is 1090:n = 1; and an ant: 1010:dosage compensation 854:, occurring in the 595:bands and sub-bands 435:staining is called 275:, is applied after 107:, and results in a 4764:Making a karyotype 3880:Systematic Biology 3266:10.1007/BF02153623 2484:. Academic Press. 2307:"Karyosystematics" 2146:Human chromosome 2 2114:hypotonic solution 2093: 2022:appearance of the 2014:, "general form") 1786: 1742: 1681:, less frequently 1668: 1558:Chromosome banding 1537:adaptive radiation 1527:All of the native 1368:Human chromosome 2 1127:Fundamental number 1108:shortnose sturgeon 824: 521: 501: 450:between the sexes, 256: 244: 158:and is designated 80: 69: 45:in the cells of a 18:Chromosome banding 5328: 5327: 5286: 5285: 5023:Centromere number 4940:Linear chromosome 4756:Media related to 4472:(10): 1063–1064. 4411:Molecular Biology 4199:978-3-8055-8019-9 4166:978-0-08-096156-9 4141:978-0-8248-2190-6 4114:978-1-4613-6877-9 3598:978-0-08-047052-8 3125:Myrmecia pilosula 3054:Muntiacus muntjak 3009:(7031): 400–404. 2127:by a solution of 2074:sex determination 1938:Angelman syndrome 1634:Silver staining: 1586:is obtained with 1458:subalpine meadows 1443:Hampton L. Carson 1385:Chilocorus stigma 1378:Some species are 1280:endoreduplication 1093:Myrmecia pilosula 1059:Muntiacus muntjak 1049:Muntiacus reevesi 1014:placental mammals 743: 742: 541:transcriptionally 317:amniotic centesis 301:white blood cells 283:by a solution of 109:photomicrographic 16:(Redirected from 5370: 5153: 5152: 5117:Polyploidization 4945:Extra chromosome 4860:Genetic material 4823: 4816: 4809: 4800: 4799: 4755: 4739: 4733: 4727: 4721: 4715: 4714: 4704: 4680: 4674: 4673: 4659: 4653: 4652: 4632: 4626: 4625: 4613: 4607: 4606: 4594: 4588: 4587: 4567: 4561: 4554: 4548: 4547: 4538:(3–4): 193–197. 4527: 4521: 4516:Delaunay L. N. 4514: 4508: 4507: 4481: 4457: 4451: 4450: 4402: 4396: 4395: 4362:(8): 1973–1986. 4356:Nature Protocols 4347: 4341: 4340: 4330: 4320: 4303:(25): 16156–61. 4288: 4282: 4281: 4237: 4231: 4230: 4210: 4204: 4203: 4185: 4179: 4178: 4152: 4146: 4145: 4125: 4119: 4118: 4092: 4086: 4085: 4069: 4063: 4062: 4040: 4034: 4033: 4023: 3987: 3981: 3980: 3955:(3938): 1414–8. 3944: 3938: 3937: 3931: 3922: 3916: 3910: 3904: 3903: 3875: 3869: 3868: 3840: 3834: 3833: 3821: 3811: 3805: 3804: 3794: 3784: 3752: 3746: 3739: 3733: 3726: 3720: 3719: 3701: 3677: 3671: 3670: 3660: 3632: 3626: 3625: 3619: 3609: 3603: 3602: 3580: 3574: 3569: 3563: 3562: 3534: 3528: 3527: 3491: 3485: 3480: 3474: 3473: 3445: 3439: 3438: 3436: 3434: 3429:on 26 March 2012 3428: 3422:. Archived from 3405: 3392: 3386: 3385: 3383: 3365: 3352: 3346: 3345: 3323: 3317: 3316: 3314: 3284: 3278: 3277: 3249: 3243: 3242: 3202: 3196: 3195: 3175: 3169: 3168: 3120: 3114: 3107: 3098: 3097: 3064:(3937): 1364–6. 3049: 3043: 3042: 2998: 2992: 2987: 2981: 2974: 2968: 2967: 2956:10.1002/jez.1131 2931: 2925: 2924: 2888: 2882: 2881: 2845: 2839: 2832: 2826: 2825: 2815: 2805: 2773: 2767: 2766: 2760: 2752: 2742: 2718: 2712: 2711: 2705: 2697: 2665: 2659: 2658: 2652: 2644: 2634: 2624: 2600: 2594: 2593: 2556:Sax, L. (2002). 2553: 2547: 2546: 2540: 2532: 2522: 2502: 2496: 2495: 2475: 2469: 2462: 2453: 2450: 2441: 2440: 2430: 2420: 2396: 2387: 2386: 2370: 2360: 2354: 2347: 2338: 2335: 2329: 2327: 2317: 2311: 2310: 2303: 2297: 2296: 2284: 2274: 2268: 2262: 2253: 2252: 2246: 2236: 2230: 2229: 2211: 2205: 2204: 2202: 2200: 2184: 2169: 2069:, concluding an 2048:Michael JD White 2044:C. D. Darlington 2032:Grigory Levitsky 1988:Walther Flemming 1890:Edwards syndrome 1572:Types of banding 1472:, in the family 1450:Hawaiian Islands 1321:palaeopolyploidy 1291:nuclear membrane 1236:, and the other 1073: 952: 872:sister chromatid 632: 631: 219:karyosystematics 93:light microscopy 21: 5378: 5377: 5373: 5372: 5371: 5369: 5368: 5367: 5333: 5332: 5329: 5324: 5282: 5189: 5151: 5121: 5110:Paleopolyploidy 5055: 5049: 4905: 4879:Heterochromatin 4842: 4836: 4827: 4794:Wayback Machine 4748: 4743: 4742: 4734: 4730: 4722: 4718: 4681: 4677: 4660: 4656: 4637:J. Exp. Zoology 4633: 4629: 4614: 4610: 4595: 4591: 4568: 4564: 4555: 4551: 4528: 4524: 4515: 4511: 4458: 4454: 4403: 4399: 4348: 4344: 4289: 4285: 4248:(5274): 494–7. 4238: 4234: 4211: 4207: 4200: 4186: 4182: 4167: 4153: 4149: 4142: 4126: 4122: 4115: 4093: 4089: 4070: 4066: 4059: 4041: 4037: 3988: 3984: 3945: 3941: 3929: 3923: 3919: 3911: 3907: 3876: 3872: 3851:(1–2): 68–169. 3841: 3837: 3830: 3812: 3808: 3753: 3749: 3740: 3736: 3727: 3723: 3678: 3674: 3651:(17): 2765–75. 3633: 3629: 3616:The chromosomes 3610: 3606: 3599: 3581: 3577: 3570: 3566: 3535: 3531: 3508:10.1038/nrg1711 3496:Nat. Rev. Genet 3492: 3488: 3481: 3477: 3446: 3442: 3432: 3430: 3426: 3403: 3393: 3389: 3363: 3353: 3349: 3324: 3320: 3285: 3281: 3250: 3246: 3203: 3199: 3176: 3172: 3121: 3117: 3109:Hsu T.C. 1979. 3108: 3101: 3050: 3046: 2999: 2995: 2988: 2984: 2975: 2971: 2932: 2928: 2889: 2885: 2846: 2842: 2833: 2829: 2774: 2770: 2754: 2753: 2719: 2715: 2699: 2698: 2666: 2662: 2646: 2645: 2601: 2597: 2554: 2550: 2534: 2533: 2503: 2499: 2492: 2478:K. Lew (2012). 2476: 2472: 2463: 2456: 2451: 2444: 2397: 2390: 2383: 2361: 2357: 2348: 2341: 2336: 2332: 2318: 2314: 2305: 2304: 2300: 2293: 2275: 2271: 2263: 2256: 2237: 2233: 2226: 2212: 2208: 2198: 2196: 2186: 2185: 2181: 2176: 2167: 2154: 2106:Using cells in 2040:Nikolai Vavilov 2036:Sergei Navashin 1998:in 1888. It is 1976: 1874:Turner syndrome 1867: 1815: 1809: 1794: 1734: 1659:probed for the 1649: 1615:Q-banding is a 1608:, so it stains 1574: 1569: 1560: 1431: 1376: 1345:Turner syndrome 1330: 1310:differentiation 1299: 1297: 1294: 1189: 1129: 1074: 1071: 1040:Kurt Benirschke 1032: 963:heterochromatin 959: 953: 950: 903: 881: 877: 863: 841: 831: 813: 615: 572:sex chromosomes 551:, and a higher 493: 491:Human karyogram 417:Giemsa staining 405:heterochromatin 329: 261: 228: 208:relationships, 183:sex chromosomes 141:sex chromosomes 35: 28: 23: 22: 15: 12: 11: 5: 5376: 5366: 5365: 5360: 5355: 5350: 5345: 5326: 5325: 5323: 5322: 5317: 5312: 5307: 5306: 5305: 5294: 5292: 5288: 5287: 5284: 5283: 5281: 5280: 5275: 5270: 5265: 5260: 5255: 5250: 5245: 5240: 5235: 5230: 5225: 5220: 5215: 5210: 5205: 5199: 5197: 5191: 5190: 5188: 5187: 5182: 5177: 5172: 5167: 5161: 5159: 5150: 5149: 5144: 5129: 5127: 5123: 5122: 5120: 5119: 5114: 5113: 5112: 5107: 5102: 5097: 5087: 5086: 5085: 5080: 5070: 5065: 5059: 5057: 5051: 5050: 5048: 5047: 5046: 5045: 5040: 5035: 5030: 5020: 5019: 5018: 5013: 5008: 5003: 5001:Submetacentric 4998: 4988: 4983: 4978: 4973: 4968: 4963: 4958: 4953: 4948: 4942: 4933: 4928: 4927:or heterosome) 4921:Sex chromosome 4913: 4911: 4907: 4906: 4904: 4903: 4898: 4893: 4888: 4883: 4882: 4881: 4876: 4866: 4857: 4852: 4846: 4844: 4838: 4837: 4826: 4825: 4818: 4811: 4803: 4797: 4796: 4784: 4779: 4773: 4767: 4761: 4747: 4746:External links 4744: 4741: 4740: 4728: 4716: 4675: 4654: 4643:(3): 291–336. 4627: 4608: 4589: 4578:(4): 465–502. 4562: 4549: 4522: 4509: 4452: 4417:(4): 542–550. 4397: 4342: 4283: 4232: 4221:(1): 229–237. 4205: 4198: 4180: 4165: 4147: 4140: 4120: 4113: 4087: 4064: 4058:978-0849365478 4057: 4035: 3982: 3939: 3917: 3905: 3886:(2): 184–194. 3870: 3835: 3828: 3806: 3767:(20): 9051–5. 3747: 3734: 3728:Nagl W. 1978. 3721: 3692:(3): 297–306. 3672: 3627: 3604: 3597: 3575: 3564: 3529: 3502:(11): 836–46. 3486: 3475: 3462:10.1086/280872 3456:(750): 54–66. 3440: 3414:(1): 113–130. 3387: 3347: 3336:(2): 131–141. 3318: 3279: 3244: 3197: 3186:(3): 205–217. 3170: 3135:(4743): 1278. 3115: 3099: 3044: 2993: 2982: 2969: 2926: 2883: 2840: 2827: 2788:(18): 9821–3. 2768: 2713: 2660: 2595: 2568:(3): 174–178. 2548: 2497: 2490: 2470: 2454: 2442: 2388: 2381: 2355: 2339: 2330: 2312: 2298: 2291: 2269: 2254: 2231: 2224: 2206: 2178: 2177: 2175: 2172: 2171: 2170: 2161: 2153: 2150: 2138: 2137: 2134: 2131: 2117: 2110: 2108:tissue culture 1975: 1972: 1971: 1970: 1951: 1945: 1935: 1929: 1919: 1918: 1915: 1909: 1906:Patau syndrome 1903: 1897: 1887: 1877: 1866: 1863: 1859:genetic mosaic 1835:nondisjunction 1823:translocations 1811:Main article: 1808: 1805: 1793: 1790: 1766:interferometer 1733: 1730: 1648: 1645: 1644: 1643: 1636:Silver nitrate 1632: 1625: 1613: 1602: 1599: 1573: 1570: 1568: 1565: 1559: 1556: 1508:moving over a 1430: 1427: 1410:Thais lapillus 1375: 1372: 1329: 1326: 1325: 1324: 1317: 1252:differentiated 1248:Endopolyploidy 1245: 1230:Haplo-diploidy 1219: 1218: 1188: 1185: 1128: 1125: 1069: 1031: 1028: 1027: 1026: 1002:X-inactivation 999: 978:Theodor Boveri 974: 958: 955: 948: 923:macromolecules 902: 899: 879: 875: 861: 839: 829: 812: 809: 808: 807: 796: 769:sex chromosome 765: 749:versus inside 741: 740: 733: 730: 724: 723: 720: 717: 711: 710: 707: 704: 698: 697: 691: 688: 682: 681: 678: 675: 669: 668: 665: 662: 656: 655: 652: 649: 643: 642: 639: 636: 614: 611: 549:non-coding DNA 492: 489: 488: 487: 480: 473: 466: 451: 441: 440: 377: 370: 366: 363: 360:translocations 352: 328: 325: 305:tissue culture 260: 257: 227: 224: 173:So, in normal 156:somatic number 26: 9: 6: 4: 3: 2: 5375: 5364: 5361: 5359: 5356: 5354: 5351: 5349: 5346: 5344: 5341: 5340: 5338: 5331: 5321: 5318: 5316: 5313: 5311: 5308: 5304: 5301: 5300: 5299: 5296: 5295: 5293: 5289: 5279: 5276: 5274: 5271: 5269: 5266: 5264: 5261: 5259: 5256: 5254: 5251: 5249: 5246: 5244: 5241: 5239: 5236: 5234: 5231: 5229: 5226: 5224: 5221: 5219: 5216: 5214: 5211: 5209: 5206: 5204: 5201: 5200: 5198: 5196: 5192: 5186: 5183: 5181: 5178: 5176: 5173: 5171: 5168: 5166: 5163: 5162: 5160: 5158: 5154: 5148: 5145: 5142: 5138: 5134: 5131: 5130: 5128: 5124: 5118: 5115: 5111: 5108: 5106: 5103: 5101: 5098: 5096: 5093: 5092: 5091: 5088: 5084: 5081: 5079: 5076: 5075: 5074: 5071: 5069: 5066: 5064: 5061: 5060: 5058: 5056:and evolution 5052: 5044: 5041: 5039: 5036: 5034: 5031: 5029: 5026: 5025: 5024: 5021: 5017: 5014: 5012: 5009: 5007: 5004: 5002: 4999: 4997: 4994: 4993: 4992: 4989: 4987: 4984: 4982: 4981:Isochromosome 4979: 4977: 4974: 4972: 4969: 4967: 4964: 4962: 4959: 4957: 4954: 4952: 4949: 4946: 4943: 4941: 4937: 4934: 4932: 4929: 4926: 4922: 4918: 4915: 4914: 4912: 4908: 4902: 4899: 4897: 4894: 4892: 4889: 4887: 4884: 4880: 4877: 4875: 4872: 4871: 4870: 4867: 4865: 4861: 4858: 4856: 4853: 4851: 4848: 4847: 4845: 4839: 4835: 4831: 4824: 4819: 4817: 4812: 4810: 4805: 4804: 4801: 4795: 4791: 4788: 4785: 4783: 4780: 4777: 4774: 4771: 4768: 4765: 4762: 4759: 4754: 4750: 4749: 4737: 4732: 4725: 4720: 4712: 4708: 4703: 4698: 4694: 4690: 4686: 4679: 4671: 4670: 4665: 4658: 4650: 4646: 4642: 4638: 4631: 4623: 4619: 4612: 4604: 4600: 4593: 4585: 4581: 4577: 4573: 4572:Bull Hist Med 4566: 4559: 4553: 4545: 4541: 4537: 4533: 4526: 4519: 4513: 4505: 4501: 4497: 4493: 4489: 4485: 4480: 4475: 4471: 4467: 4463: 4456: 4448: 4444: 4440: 4436: 4432: 4428: 4424: 4420: 4416: 4412: 4408: 4401: 4393: 4389: 4385: 4381: 4377: 4373: 4369: 4365: 4361: 4357: 4353: 4346: 4338: 4334: 4329: 4324: 4319: 4314: 4310: 4306: 4302: 4298: 4294: 4287: 4279: 4275: 4271: 4267: 4263: 4259: 4255: 4251: 4247: 4243: 4236: 4228: 4224: 4220: 4216: 4209: 4201: 4195: 4191: 4184: 4176: 4172: 4168: 4162: 4158: 4151: 4143: 4137: 4133: 4132: 4124: 4116: 4110: 4106: 4102: 4098: 4091: 4083: 4079: 4075: 4068: 4060: 4054: 4050: 4046: 4039: 4031: 4027: 4022: 4017: 4013: 4009: 4006:(3): 465–82. 4005: 4001: 3997: 3995: 3986: 3978: 3974: 3970: 3966: 3962: 3958: 3954: 3950: 3943: 3935: 3928: 3921: 3915:, p. 169 3914: 3909: 3901: 3897: 3893: 3889: 3885: 3881: 3874: 3866: 3862: 3858: 3854: 3850: 3846: 3839: 3831: 3829:9780387076683 3825: 3820: 3819: 3810: 3802: 3798: 3793: 3788: 3783: 3778: 3774: 3770: 3766: 3762: 3758: 3751: 3744: 3738: 3731: 3725: 3717: 3713: 3709: 3705: 3700: 3695: 3691: 3687: 3683: 3676: 3668: 3664: 3659: 3654: 3650: 3646: 3642: 3640: 3631: 3623: 3618: 3617: 3608: 3600: 3594: 3590: 3586: 3579: 3573: 3572:Stebbins 1971 3568: 3560: 3556: 3552: 3548: 3545:(2): 135–41. 3544: 3540: 3533: 3525: 3521: 3517: 3513: 3509: 3505: 3501: 3497: 3490: 3484: 3483:Stebbins 1950 3479: 3471: 3467: 3463: 3459: 3455: 3451: 3444: 3425: 3421: 3417: 3413: 3409: 3402: 3400: 3391: 3382: 3377: 3373: 3369: 3362: 3360: 3351: 3343: 3339: 3335: 3331: 3330: 3322: 3313: 3308: 3304: 3300: 3299: 3294: 3292: 3283: 3275: 3271: 3267: 3263: 3259: 3255: 3248: 3240: 3236: 3232: 3228: 3224: 3220: 3216: 3212: 3208: 3201: 3193: 3189: 3185: 3181: 3174: 3166: 3162: 3158: 3154: 3150: 3146: 3142: 3138: 3134: 3130: 3126: 3119: 3112: 3106: 3104: 3095: 3091: 3087: 3083: 3079: 3075: 3071: 3067: 3063: 3059: 3055: 3048: 3040: 3036: 3032: 3028: 3024: 3020: 3016: 3012: 3008: 3004: 2997: 2991: 2986: 2979: 2973: 2965: 2961: 2957: 2953: 2949: 2945: 2941: 2937: 2930: 2922: 2918: 2914: 2910: 2906: 2902: 2898: 2894: 2887: 2879: 2875: 2871: 2867: 2863: 2859: 2856:(3): 242–50. 2855: 2851: 2844: 2837: 2831: 2823: 2819: 2814: 2809: 2804: 2799: 2795: 2791: 2787: 2783: 2779: 2772: 2764: 2758: 2750: 2746: 2741: 2736: 2732: 2728: 2724: 2717: 2709: 2703: 2695: 2691: 2687: 2683: 2680:(3): 1360–5. 2679: 2675: 2671: 2664: 2656: 2650: 2642: 2638: 2633: 2628: 2623: 2618: 2614: 2610: 2606: 2599: 2591: 2587: 2583: 2579: 2575: 2571: 2567: 2563: 2559: 2552: 2544: 2538: 2530: 2526: 2521: 2516: 2512: 2508: 2501: 2493: 2491:9780123813749 2487: 2483: 2482: 2474: 2467: 2461: 2459: 2449: 2447: 2438: 2434: 2429: 2424: 2419: 2414: 2410: 2406: 2402: 2395: 2393: 2384: 2382:9780713122879 2378: 2374: 2369: 2368: 2359: 2352: 2346: 2344: 2334: 2325: 2324: 2316: 2308: 2302: 2294: 2292:9780231017336 2288: 2283: 2282: 2273: 2266: 2261: 2259: 2250: 2245: 2244: 2235: 2227: 2225:0-87893-403-0 2221: 2217: 2210: 2195: 2194: 2189: 2183: 2179: 2165: 2164:Genome screen 2162: 2159: 2156: 2155: 2149: 2147: 2143: 2135: 2132: 2130: 2126: 2122: 2118: 2115: 2111: 2109: 2105: 2104: 2103: 2101: 2097: 2089: 2085: 2083: 2079: 2075: 2072: 2068: 2064: 2063:spermatogonia 2060: 2056: 2051: 2049: 2045: 2041: 2037: 2033: 2029: 2025: 2021: 2015: 2013: 2009: 2005: 2004:Ancient Greek 2001: 1997: 1993: 1989: 1985: 1981: 1968: 1964: 1960: 1956: 1952: 1949: 1946: 1943: 1939: 1936: 1933: 1930: 1927: 1924: 1923: 1922: 1916: 1913: 1910: 1907: 1904: 1901: 1900:Down syndrome 1898: 1895: 1892:is caused by 1891: 1888: 1885: 1881: 1878: 1875: 1872: 1871: 1870: 1862: 1860: 1856: 1852: 1848: 1844: 1840: 1836: 1832: 1828: 1824: 1820: 1814: 1804: 1802: 1798: 1789: 1782: 1778: 1776: 1771: 1767: 1763: 1759: 1758:Fluorescently 1755: 1751: 1747: 1738: 1729: 1727: 1723: 1719: 1715: 1711: 1707: 1702: 1700: 1696: 1692: 1688: 1684: 1680: 1677: 1673: 1666: 1662: 1658: 1653: 1641: 1637: 1633: 1630: 1626: 1622: 1618: 1614: 1611: 1607: 1603: 1600: 1597: 1593: 1589: 1585: 1582: 1581: 1580: 1578: 1564: 1555: 1552: 1550: 1546: 1542: 1538: 1534: 1530: 1525: 1523: 1519: 1515: 1511: 1507: 1506:Pacific Plate 1503: 1499: 1495: 1491: 1485: 1483: 1477: 1475: 1474:Drosophilidae 1471: 1470: 1465: 1464: 1459: 1455: 1451: 1446: 1444: 1440: 1436: 1429:Species trees 1426: 1424: 1420: 1416: 1412: 1411: 1407: 1403: 1402: 1401:Sorex araneus 1397: 1396: 1392:of the genus 1391: 1387: 1386: 1381: 1371: 1369: 1365: 1361: 1360: 1355: 1354: 1348: 1346: 1342: 1341:Down syndrome 1338: 1334: 1322: 1318: 1316:in many ways. 1315: 1314:morphogenesis 1311: 1307: 1304:kingdom from 1303: 1292: 1289: 1285: 1284:reduplication 1281: 1277: 1273: 1269: 1265: 1261: 1257: 1253: 1249: 1246: 1243: 1239: 1235: 1231: 1228: 1227: 1226: 1224: 1216: 1212: 1208: 1204: 1200: 1197: 1196: 1195: 1193: 1184: 1182: 1178: 1174: 1170: 1166: 1162: 1158: 1154: 1150: 1146: 1142: 1138: 1134: 1124: 1122: 1118: 1117:B chromosomes 1114: 1113: 1109: 1105: 1104: 1099: 1095: 1094: 1089: 1085: 1084: 1080: 1068: 1063: 1061: 1060: 1055: 1051: 1050: 1045: 1044:Doris Wurster 1041: 1037: 1024: 1019: 1015: 1011: 1007: 1003: 1000: 997: 993: 992: 987: 983: 979: 975: 972: 971:sciarid flies 968: 967: 966: 964: 951:Maynard Smith 947: 943: 938: 933: 930: 928: 924: 920: 916: 912: 911:transcription 908: 905:Although the 898: 896: 892: 887: 885: 873: 869: 865: 857: 853: 852:DNA synthesis 849: 845: 842:phase of the 837: 833: 822: 817: 805: 801: 797: 794: 790: 786: 782: 781:X chromosomes 778: 774: 770: 766: 763: 760: 756: 755: 754: 752: 748: 738: 734: 731: 729: 726: 725: 721: 718: 716: 713: 712: 708: 705: 703: 700: 699: 696: 692: 689: 687: 684: 683: 679: 676: 674: 671: 670: 666: 663: 661: 658: 657: 653: 650: 648: 645: 644: 640: 637: 634: 633: 630: 628: 624: 620: 610: 608: 604: 600: 596: 592: 587: 585: 581: 577: 573: 569: 565: 561: 556: 554: 550: 546: 542: 538: 534: 530: 526: 518: 514: 510: 505: 497: 485: 481: 478: 474: 471: 467: 464: 460: 456: 452: 449: 448: 447: 444: 438: 434: 430: 426: 422: 418: 414: 410: 406: 402: 398: 394: 391:pairs versus 390: 386: 382: 378: 375: 371: 367: 364: 361: 357: 353: 350: 346: 345: 340: 339: 334: 333: 332: 324: 322: 318: 314: 310: 306: 302: 298: 294: 290: 286: 282: 281:cell division 278: 274: 270: 266: 253: 248: 241: 237: 232: 223: 221: 220: 215: 211: 207: 203: 199: 194: 192: 188: 184: 180: 176: 171: 169: 165: 161: 157: 153: 148: 146: 142: 138: 134: 130: 125: 123: 118: 114: 110: 106: 102: 98: 94: 89: 85: 77: 73: 66: 62: 58: 54: 52: 48: 44: 40: 33: 19: 5353:Cytogenetics 5343:Cell biology 5330: 5022: 4990: 4849: 4830:Cytogenetics 4738:(video) 2007 4731: 4719: 4695:(1–2): 1–6. 4692: 4688: 4678: 4669:The Guardian 4667: 4657: 4640: 4636: 4630: 4621: 4617: 4611: 4605:(93): 147–9. 4602: 4598: 4592: 4575: 4571: 4565: 4557: 4552: 4535: 4531: 4525: 4517: 4512: 4469: 4465: 4455: 4414: 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