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.

493: 1601:. 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. 1641: 46: 485: 61: 805: 1770: 1726: 4742: 863:(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 1792:. 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. 236: 1753:. 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 925:

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

1583:. 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, 1903:, 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. 1761:

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

1206:) 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. 1055:

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

1351:, 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 983:, 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 1590:

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

910:, 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 79:

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.

1124:, 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 ( 388:). 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 1371:

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.

1194:. 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 ( 2069:

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

2546: 1190:, 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 4084:

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.

1838:, 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 5308: 587: 3385: 1690:-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. 575:, 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). 239:

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.

839:, 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 1713:

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

1917:(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. 512:

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.

227:. Karyograms are generally made by chromosomes in prometaphase or metaphase. During these phases, the two copies of each chromosome (connected at the 1842:. 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 2751: 2696: 2643: 2531: 1693:

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

1032: 827:) 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 4809: 1984: 1486: 4752: 2923:

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

1939:– 50% of cases have a segment of the long arm of chromosome 15 missing; a deletion of the paternal genes, example of imprinting disorder. 4770: 1543:

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".

795:), although this is not included in micrographic karyograms in clinical practice. Its genome is relatively tiny compared to the rest. 1066:

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

859:(annotated as "Meta."). During this interval, there is still 2n, but each chromosome will have 2 copies of each locus, wherein each 5206: 1631:-associated protein. This yields a dark region where the silver is deposited, denoting the activity of rRNA genes within the NOR. 1324:

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".

1005:, the inactivation is random as between the two Xs; thus the mammalian female is a mosaic in respect of her X chromosomes. In 5298: 4939: 4186: 4153: 4128: 4101: 3585: 2042:

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".

4045: 1734: 2496:"Relevance of human chromosome analysis activities against mutation concept in genetics course. IOP Conference Series" 4802: 3816: 3412: 2478: 2369: 2279: 2212: 1929:– 50% of cases have a segment of the long arm of chromosome 15 missing; a deletion of the maternal genes, example of 1653: 1446: 4229:

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

1701:, respectively. In addition, the differently stained regions and sub-regions are given numerical designations from 1426:

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

760:(in green rectangle at bottom right in the schematic karyogram, with adjacent silhouettes of typical representative 5078: 4758: 3194:

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).

835:), each cell has 2 autosomal chromosomes of each kind (designated 2n), where each chromosome has one copy of each 4764: 4652: 1510: 104:

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"

3278:"Karyotype characterization and ZZ/ZW sex chromosome heteromorphism in two species of the catfish genus 4063:"Chromosomes and male genitalia of Hawaiian Drosophila: tools for interpreting phylogeny and geography" 3317: 2181: 1298: 1968: 5071: 3980: 1811: 2176: 5125: 4909: 2312:

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

2062: 1839: 1801: 1240: 899: 791:(shown at bottom left in the schematic karyogram, to scale compared to the nuclear DNA in terms of 2470:

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

3286: 1947: 529: 993:. The inactivation of one X chromosome takes place during the early development of mammals (see 5031: 5004: 4549:. Cambridge University Press. 2nd ed, revised and enlarged, 1958. Oliver & Boyd, Edinburgh. 2361: 2237: 1920: 1749:

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.

5341: 5331: 5066: 4964: 4070: 2745: 2690: 2637: 2525: 1936: 1871:, the most common male chromosomal disease, otherwise known as 47,XXY, is caused by an extra 1815: 1807: 1529: 1470: 1149: 1145: 1141: 1137: 1133: 747: 611: 465: 132:, and any other physical characteristics. The preparation and study of karyotypes is part of 2342:

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).

4293: 4238: 3945: 3757: 3207: 3125: 3054: 2999: 2932: 2778: 2591: 2310: 2146: 1868: 1071: 725: 683: 492: 362: 1132:) present. Humans have FN = 82, due to the presence of five acrocentric chromosome pairs: 950:

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,

4690: 4673: 4488: 4431: 4376: 4262: 4009: 3845: 3700: 3508: 3454: 3258: 3223: 3180: 3149: 3078: 3023: 2905: 2862: 2728: 2711: 2620: 2593: 2574: 2513: 2416: 2389: 2354: 2268: 2134: 2091:'s lab found the chromosome count to be 46 using new techniques available at the time: 2066: 1930: 1525: 1498: 1478: 1356: 1096: 521: 496:

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.

2012: 1926: 1789: 1640: 1449:. These roughly 800 Hawaiian drosophilid species are usually assigned to two genera, 1373: 1268: 1081: 1047: 1037: 1002: 472: 447: 289: 231:) will look as one unless the image resolution is high enough to distinguish the two. 140: 4492: 4380: 4266: 3704: 3458: 3369: 3300: 3277: 3262: 3227: 3153: 3082: 2909: 2880:

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

5105: 4984: 4755:, an online activity from the University of Utah's Genetic Science Learning Center. 4685: 4633: 4532: 4528: 4519:

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).

3343: 3326: 3295: 3250: 3215: 3176: 3133: 3062: 3027: 3007: 2940: 2889: 2846: 2796: 2786: 2723: 2670: 2615: 2605: 2558: 2503: 2411: 2401: 2036: 2032: 2020: 1976: 1878: 1714: 1438: 1345:, where the gametic (= haploid) numbers form the series x = 3, 4, 5, 6, and 7; and 1279: 1129: 1125: 860: 836: 583: 81: 3957: 3137: 3066: 2122:

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".

2102: 1818:, large-scale deletions or duplications. Numerical abnormalities, also known as 1441:

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.

1894: 1847: 1823: 1754: 1624: 1398: 1236: 990: 966: 757: 567:(sperm and egg cells) which are haploid with 23 unpaired chromosomes, and this 537: 348: 320:

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

1164:, of a karyotype is the number of visible major chromosomal arms per set of 1023:

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.

4944: 4818: 4657: 4637: 4484: 4427: 4372: 4356: 4325: 4306: 4215: 4018: 3696: 3655: 3646: 3625: 3547: 3504: 3145: 3019: 2952: 2893: 2858: 2810: 2737: 2629: 2570: 2425: 2088: 2084: 1649: 1584: 1576: 1565: 1533: 1252: 1248: 1153: 1091: 1045:. When they looked at the karyotype of the closely related Indian muntjac, 979: 915: 914:. This variation provides the basis for a range of studies in evolutionary 777: 769: 739: 735: 525: 497: 326: 281: 235: 202: 190: 133: 4572: 4258: 4060: 3965: 3853: 3789: 3167:

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

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employs several techniques to visualize different aspects of chromosomes:

60: 5163: 5158: 5021: 4999: 4994: 4862: 4699: 2493: 1914: 1763: 1758: 1750: 1746: 1738: 1725: 1710: 1605: 1264: 1256: 1230: 1042: 389: 344: 207: 85: 31: 3011: 2767:"Kinetochore reproduction theory may explain rapid chromosome evolution" 2031:. The subsequent history of the concept can be followed in the works of 1674:, is used to stain bands on the chromosomes. Giemsa is specific for the 1493:, which can be dated to 30 mya. The archipelago itself (produced by the 1489:). The oldest member of the Hawaiian archipelago still above the sea is 1051:, they were astonished to find it had female = 6, male = 7 chromosomes. 5183: 5173: 5168: 5153: 5093: 5083: 4959: 4933: 4884: 4874: 4822: 3350:(Rodentia: Sigmodontinae) from Chapada Diamantina, northeastern Brazil" 3254: 2130: 2117: 2008: 1972: 1819: 1742: 1645: 1609: 1598: 1502: 1490: 1457: 1451: 1411: 1352: 1321: 1203: 1199: 1187: 1011: 1006: 974: 907: 832: 824: 809: 804: 607: 595: 579: 555:

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

2207:(2 ed.). Sunderland MA, US: Sinauer Associates Inc. p. 544. 1891:, a common chromosomal disease, is caused by trisomy of chromosome 21. 1359:

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

3746:"Origin of human chromosome 2: an ancestral telomere-telomere fusion" 2944: 2709: 2656: 2390:"Analytical Biases Associated with GC-Content in Molecular Evolution" 2113: 1988: 1900: 1675: 1671: 1572: 1482: 1403: 1290: 1109: 994: 959: 911: 872: 856: 792: 761: 750: 572: 547:

Both the micrographic and schematic karyograms show the normal human

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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) (

1407: 1294: 1169: 1165: 1067: 970: 883: 848: 816: 781: 780:

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

618:), the human chromosomes are classified into the following groups: 556: 413: 377: 253: 198: 20: 4586:

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

3802: 3744:

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

516:, which is the appearance of the chromosomes after treatment with 5291: 5145: 5056: 5051: 4775: 4280:

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

3981:"Chromosomal sequences and interisland colonizations in Hawaiian 3916:"The Hawaiian-Emperor volcanic chain, Part I. Geologic evolution" 2109: 2055: 2043: 1980: 1882: 1858:

Chromosomal abnormalities that lead to disease in humans include

1843: 1835: 1827: 1687: 1683: 1580: 1394: 1378: 1244: 1226: 1222: 1211: 1076: 1024: 1014:

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

844: 765: 548: 517: 451: 409: 385: 381: 373: 224: 170:

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

163: 35: 4624:

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

3807:

A glossary of genetics and cytogenetics: Classical and molecular

2594:"Measuring DNA content in live cells by fluorescence microscopy" 1275:, the daughter chromosomes separating from each other inside an 606:

Based on the karyogram characteristics of size, position of the

5201: 5088: 4852: 4848: 4843: 4741: 3193: 1943: 1831: 1664: 1537: 1383: 1347: 1341: 1180: 987:, all the somatic cell precursors undergo chromatin diminution. 847:(annotated as S) of the cell cycle. This interval includes the 568: 564: 552: 421: 285: 261: 179: 53: 4393: 3282:

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

1721:

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

500:, with numbered chromosome pairs, its main changes during the 5266: 5251: 5246: 5241: 5236: 5231: 5226: 5221: 5216: 5191: 5129: 4176: 3198:

with the highest chromosome number in the Acipenseriformes".

1741:

techniques used to simultaneously visualize all the pairs of

1717:

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

1506: 1473:, make it possible to see which species are closely related. 882:

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

773: 508:

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

301: 16:

Photographic display of total chromosome complement in a cell

4713:

Human chromosome 2 is a fusion of two ancestral. chromosomes

3111: 1983:, in 1882. The name was coined by another German anatomist, 886:), mainly depending on the number of mitochondria per cell. 3913: 2016: 1195: 1086: 582:, and the scale to the right shows the designations of the 365:

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

185:

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

4338: 4067:

Hawaiian biogeography: evolution on a hot spot archipelago

3100:

Human and mammalian cytogenetics: a historical perspective

2202: 4069:. Washington DC: Smithsonian Institution Press. pp. 1923:, from the loss of part of the short arm of chromosome 1. 1679: 1660: 1422:

The detailed study of chromosome banding in insects with

1183:

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

954:, or other kinds of visible adjustment to the karyotype. 903: 304:

can be predicted by observation of interphase cells (see

257: 124:. Attention is paid to their length, the position of the 3921:. In Decker, R.W.; Wright, T.L.; Stauffer, P.H. (eds.). 1850:

individual who has some normal and some abnormal cells.

1312:

for the investigation of ancient karyotype duplications.

889: 563:(allosomes). A major exception to diploidy in humans is 5309:

International System for Human Cytogenetic Nomenclature

4653:"Joe Hin Tjio The man who cracked the chromosome count" 4228: 4061:

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

3041:

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

2157:

Pages displaying short descriptions of redirect targets

962:, entire chromosomes are eliminated during development. 588:

International System for Human Cytogenomic Nomenclature

4279: 4201: 3743: 3623: 3383: 2978: 2227: 2149: – Symbols and abbreviations used in cytogenetics 1104:

at 372 chromosomes. The existence of supernumerary or

296:. Sometimes observations may be made on non-dividing ( 2712:"Mitochondria in oocyte aging: current understanding" 2266:

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

2228:

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

1437:

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

3609:(6th ed.). London: Chapman & Hall. p. 2969:. Sinauer Associates, Stamford CT. 8th ed, Chapter 9 2334: 2332: 1950:, a translocation mutation commonly associated with 958:

Chromosome elimination. In some species, as in many

926:

related species, which were previously inexplicable.

4605:Painter T.S. (1922). "The spermatogenesis of man". 2879: 2441:

Thompson & Thompson Genetics in Medicine 7th Ed

2387: 1865:

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

594:. Each row of chromosomes is vertically aligned at 551:karyotype, which is the typical composition of the 361:Differences in number and position of satellites. 120:and what these chromosomes look like under a light 100:(or simply micrographic) karyogram. In contrast, a 19:"Idiogram" redirects here. Not to be confused with 3804: 3602: 2659:"Mitochondrial DNA in anucleate human blood cells" 2494:Erwinsyah, R., Riandi, & Nurjhani, M. (2017). 2353: 2267: 1967:Chromosomes were first observed in plant cells by 1587:. It is the most common chromosome banding method. 1289:The phenomenon occurs sporadically throughout the 823:of the cellular state (outside of the replicative 143:cells of an individual or a species is called the 4761:from the University of Arizona's Biology Project. 4671: 4585: 3040: 2329: 2236:(7th ed.). Oxford University Press. p. 1737:and the older spectral karyotyping are molecular 1635: 1297:to humans; it is diverse and complex, and serves 1085:. The high record would be somewhere amongst the 1018: 784:, sometimes due to variations in sex chromosomes. 347:. These differences probably came about through 30:is the general appearance of the complete set of 5323: 4120:Hawaiian natural history, ecology, and evolution 3045:: a deer with a low diploid chromosome number". 2383: 2381: 1501:) has existed for far longer, at least into the 1355:have 24x2 chromosomes whereas humans have 23x2. 408:. Euchromatin regions contain larger amounts of 4725:Evidence of common ancestry: human chromosome 2 3803:Rieger, R.; Michaelis, A.; Green, M.M. (1968). 3668: 3626:"New insights into cell cycle control from the 3384:Weksler, M.; Bonvicino, C.R. (3 January 2005). 2922: 2105:, which swells them and spreads the chromosomes 1962: 128:, banding pattern, any differences between the 4449:Vermeesch, Joris Robert; Rauch, Anita (2006). 4448: 4177:Lisa G. Shaffer; Niels Tommerup, eds. (2005). 4040:. Boca Raton FL: CRC Press. pp. 407–439. 2764: 2710:Zhang D, Keilty D, Zhang ZF, Chian RC (2017). 2657:Shuster RC, Rubenstein AJ, Wallace DC (1988). 1393:. There is some evidence from the case of the 178:cells have multiple copies of chromosomes and 4817: 4803: 4083: 3166: 3116:, an ant with only one pair of chromosomes". 2989: 2437: 2435: 2378: 2308: 1536:. Although it would be possible for a single 1271:) chromosomes in a 'resting' nucleus undergo 1035:. The diploid number of the Chinese muntjac, 945: 591: 428:and therefore produces the typical "G-Bands". 214: 4623: 4604: 4032:Carson H.L. (1992). "Inversions in Hawaiian 3671:"Endoreplication cell cycles: more for less" 3571: 2836: 2750:: CS1 maint: multiple names: authors list ( 2703: 2695:: CS1 maint: multiple names: authors list ( 2642:: CS1 maint: multiple names: authors list ( 2585: 2530:: CS1 maint: multiple names: authors list ( 1999:, "kernel", "seed", or "nucleus", and τύπος 1942:Chromosomal abnormalities can also occur in 252:The study of karyotypes is made possible by 4396:"Genome: Origins and evolution of the term" 4143: 4031: 3525: 3308: 2650: 2338:Gustashaw K.M. 1991. Chromosome stains. In 1795: 1362: 1243:tissues the cells have ceased to divide by 586:. Such bands and sub-bands are used by the 159:(humans: n = 23). Thus, in humans 2n = 46. 4810: 4796: 4771:Chromosome Staining and Banding Techniques 4036:". In Krimbas, C.B.; Powell, J.R. (eds.). 3831: 3397:Arquivos do Museu Nacional, Rio de Janeiro 2432: 2205:Plant systematics, a phylogenetic approach 1555: 1410:coast, that the two chromosome morphs are 601: 520:(to partially digest the chromosomes) and 368:Differences in degree and distribution of 4776:Bjorn Biosystems for Karyotyping and FISH 4689: 4518: 4466: 4315: 4305: 4144:Maloy, Stanley R.; Hughes, Kelly (2013). 4008: 3978: 3935: 3779: 3769: 3686: 3645: 3368: 3299: 2800: 2790: 2727: 2619: 2609: 2507: 2449: 2447: 2415: 2405: 1659:In the "classic" (depicted) karyotype, a 724:Very small, acrocentric (and 21, 22 with 155:(the sex cells) the chromosome number is 4759:Karyotyping activity with case histories 3560: 3471: 3436: 3269: 3169:Botanical Journal of the Linnean Society 3112:Crosland M.W.J.; Crozier, R.H. (1986). " 2351: 2265: 2249: 2247: 2075: 1768: 1724: 1639: 1579:following digestion of chromosomes with 1505:. Previous islands now beneath the sea ( 803: 491: 483: 336:each have six pairs of chromosomes, yet 234: 218: 59: 44: 4116: 3482: 3377: 3240: 3234: 3187: 965:Chromatin diminution (founding father: 139:The basic number of chromosomes in the 5324: 4706: 4650: 3914:Clague, D.A.; Dalrymple, G.B. (1987). 3866: 3732:Chromosomal evolution in higher plants 3094: 3092: 2765:Godfrey LR, Masters JC (August 2000). 2466: 2444: 2356:Chromosomal evolution in higher plants 2340:The ACT Cytogenetics Laboratory Manual 1897:is caused by trisomy of chromosome 13. 1780: 1709:on the chromosome arms. For example, 284:when most condensed. In order for the 5299:List of organisms by chromosome count 4791: 4715:Alec MacAndrew; accessed 18 May 2006. 3901: 3617: 3600: 3337: 2457:. 3rd ed, Cambridge University Press. 2253: 2244: 1546: 1115: 890:Diversity and evolution of karyotypes 698:Small, metacentric or submetacentric 643:Large, metacentric or submetacentric 201:and to gather information about past 4065:. In Wagner, W.L.; Funk, E. (eds.). 3669:Edgar BA, Orr-Weaver TL (May 2001). 3580:. Academic Press. pp. 427–517. 2979:King, Stansfield & Mulligan 2006 1745:in an organism in different colors. 1729:Spectral karyogram of a human female 1229:. It is a common arrangement in the 871:phases, nuclear DNA is dispersed as 396:. GC rich DNA tends to contain more 4651:Wright, Pearce (11 December 2001). 3572:Gregory, T.R.; Mable, B.K. (2011). 3386:"Taxonomy of pygmy rice rats genus 3089: 2544: 2196: 1971:in 1842. Their behavior in animal ( 1560: 13: 4691:10.1111/j.1601-5223.1956.tb03010.x 4558: 4455:European Journal of Human Genetics 4146:Brenner's Encyclopedia of Genetics 3846:10.1111/j.1601-5223.1969.tb02259.x 3181:10.1111/j.1095-8339.1990.tb01876.x 2015:chromosomes, in contrast to their 969:). In this process, found in some 764:): The most common karyotypes for 479: 475:or otherwise abnormal individuals. 14: 5363: 4734: 4148:. San Diego, CA: Academic Press. 4038:Drosophila inversion polymorphism 3624:Lilly M.A.; Duronio R.J. (2005). 2500:Materials Science and Engineering 2317:Revised August 2004, January 2008 2270:Variation and evolution in plants 457:between members of a population ( 223:Chromosomes at various stages of 4740: 1885:(three copies) of chromosome 18. 1417: 1285:division' is not quite accurate. 682:Medium-sized, acrocentric, with 610:and sometimes the presence of a 532:active, with a greater ratio of 420:). The staining technique using 416:pairs (that is, it has a higher 4920:Macrochromosome/Microchromosome 4718: 4665: 4644: 4617: 4598: 4579: 4552: 4539: 4512: 4499: 4442: 4387: 4332: 4273: 4222: 4195: 4170: 4137: 4110: 4088:. Vol. 31. pp. 1–43. 4077: 4054: 4025: 3972: 3929: 3907: 3895: 3860: 3825: 3796: 3737: 3730:Stebbins, G. Ledley, Jr. 1972. 3724: 3711: 3662: 3594: 3565: 3554: 3519: 3476: 3465: 3430: 3370:10.1590/S1984-46702011000100013 3301:10.1590/S1679-62252007000300010 3160: 3105: 3034: 2983: 2972: 2959: 2916: 2873: 2830: 2817: 2758: 2538: 2487: 2460: 2345: 1251:contain more than the original 1089:, with the adder's tongue fern 343:Differences in the position of 315: 118:chromosome count of an organism 4674:"The chromosome number of man" 4533:10.1080/00087114.1994.10797297 4123:. University of Hawaii Press. 3574:"Ch. 8: Polyploidy in animals" 2509:10.1088/1757-899x/180/1/012285 2320: 2302: 2288: 2259: 2221: 2169: 2023:who worked with Lev Delaunay, 1644:Karyogram from a human female 1636:Classic karyotype cytogenetics 1019:Number of chromosomes in a set 799: 300:) cells. The sex of an unborn 52:karyogram of human male using 1: 4451:"Reply to Hochstenbach et al" 4181:. Switzerland: S. Karger AG. 3958:10.1126/science.168.3938.1414 3811:. New York: Springer-Verlag. 3688:10.1016/S0092-8674(01)00334-8 3576:. In Gregory, T. Ryan (ed.). 3331:10.1016/S0047-2484(82)80045-6 3138:10.1126/science.231.4743.1278 3067:10.1126/science.168.3937.1364 2675:10.1016/s0006-291x(88)81291-9 2455:Animal cytology and evolution 2274:. Columbia University Press. 2162: 2046:human cell contain? In 1912, 1822:, often occur as a result of 1629:nucleolar organization region 1316: 776:usually have both an X and a 669:Medium-sized, submetacentric 4672:Tjio J.H.; Levan A. (1956). 4547:Evolution of genetic systems 4286:Proc. Natl. Acad. Sci. U.S.A 4251:10.1126/science.273.5274.494 3750:Proc. Natl. Acad. Sci. U.S.A 2771:Proc. Natl. Acad. Sci. U.S.A 2388:Romiguier J, Roux C (2017). 1963:History of karyotype studies 1956:acute lymphoblastic leukemia 1952:chronic myelogenous leukemia 1853: 1595:constitutive heterochromatin 1027:, which was investigated by 559:chromosomes and one pair of 404:. GC rich DNA is lighter on 7: 4765:Printable karyotype project 4094:10.1007/978-1-4615-4185-1_1 3578:The Evolution of the Genome 2360:. London: Arnold. pp. 2140: 2050:reported 47 chromosomes in 2019:contents was introduced by 1593:C-banding: Giemsa binds to 1469:rearrangements, especially 1233:, and in some other groups. 247: 10: 5368: 4960:Dinoflagellate chromosomes 4001:10.1093/genetics/103.3.465 3318:Journal of Human Evolution 2663:Biochem Biophys Res Commun 2315:. Oxford University Press. 2182:Collins English Dictionary 2155: – Laboratory process 1799: 1682:. Quinacrine binds to the 1041:, was found to be 46, all 946:Changes during development 906:is highly standardized in 880:human mitochondrial genome 789:human mitochondrial genome 454:and the rest of the body), 435:Variation is often found: 268:have been arrested during 215:Observations on karyotypes 182:cells have single copies. 18: 5304:List of sequenced genomes 5279: 5182: 5144: 5114: 5072:Chromosomal translocation 5042: 4945:A chromosome/B chromosome 4936:(or accessory chromosome) 4898: 4829: 4412:10.1134/S0026893316040178 4117:Ziegler, Alan C. (2002). 3540:10.1016/j.pbi.2005.01.001 3220:10.1007/s10228-004-0257-z 2611:10.1186/s13008-018-0039-z 2563:10.1080/00224490209552139 2547:"How Common is Intersex?" 1975:) cells was described by 1528:was spurred by a lack of 1175: 590:to describe locations of 466:geographic specialization 5126:Telomere-binding protein 4940:Supernumerary chromosome 3979:Carson HL (March 1983). 2827:. 2nd ed, Oxford. p218-9 2407:10.3389/fgene.2017.00016 2232:A dictionary of genetics 1840:homologous recombination 1802:Chromosome abnormalities 1796:Chromosome abnormalities 1773:Spectral human karyotype 1363:Chromosomal polymorphism 1120:The fundamental number, 711:Very small, metacentric 592:chromosome abnormalities 402:transcriptionally active 116:Karyotypes describe the 4468:10.1038/sj.ejhg.5201663 3881:10.1093/sysbio/33.2.184 3771:10.1073/pnas.88.20.9051 3439:The American Naturalist 3287:Neotropical Ichthyology 3200:Ichthyological Research 2823:Maynard Smith J. 1998. 2792:10.1073/pnas.97.18.9821 2551:Journal of Sex Research 2352:Stebbins, G.L. (1971). 2177:"Karyotype, definition" 2101:Pretreating cells in a 1969:Carl Wilhelm von Nägeli 1948:Philadelphia chromosome 1608:pattern obtained using 1556:Depiction of karyotypes 1414:to different habitats. 878:The copy number of the 772:and are denoted 46,XX; 614:(a segment distal to a 602:Human chromosome groups 459:chromosome polymorphism 187:chromosomal aberrations 5062:Structural alterations 4638:10.1002/jez.1400370303 4545:Darlington C.D. 1939. 4357:10.1038/nprot.2007.276 4307:10.1073/pnas.202610899 3647:10.1038/sj.onc.1208610 3601:White, M.J.D. (1973). 3528:Curr. Opin. Plant Biol 3196:Acipenser brevirostrum 3102:. Springer-Verlag, NY. 2894:10.1002/bies.950180209 2309:Lee M. Silver (1995). 2081: 1921:1p36 Deletion syndrome 1830:in the formation of a 1774: 1730: 1672:mepacrine (quinacrine) 1656: 1532:and a wide variety of 1511:Emperor Seamount Chain 1336:are examples of this. 1326:chromosome abnormality 1170:sex-linked chromosomes 1101:Acipenser brevirostrum 1064: 943: 928: 812: 656:Large, submetacentric 616:secondary constriction 509: 504:(top center), and the 489: 256:. Usually, a suitable 244: 232: 68: 57: 5079:Numerical alterations 5067:Chromosomal inversion 4965:Homologous chromosome 4341:"Digital karyotyping" 4282:"Digital karyotyping" 3734:. Nelson, London. p18 3721:. Elsevier, New York. 3348:Wiedomys pyrrhorhinus 2967:Developmental biology 2825:Evolutionary genetics 2716:Facts Views Vis Obgyn 2133:have 48 chromosomes. 2079: 1985:Heinrich von Waldeyer 1937:Prader-Willi syndrome 1808:derivative chromosome 1772: 1728: 1643: 1616:T-banding: visualize 1428:Hawaiian drosophilids 1387:, the European shrew 1239:occurs when in adult 1053: 933: 923: 807: 612:chromosomal satellite 495: 487: 238: 222: 63: 48: 5347:Evolutionary biology 5287:Extrachromosomal DNA 4975:Satellite chromosome 4950:Lampbrush chromosome 4890:Nuclear organization 4781:12 June 2019 at the 4749:at Wikimedia Commons 4588:Archives de Biologie 4086:Evolutionary Biology 3346:"A new karyotype of 2147:Cytogenetic notation 1979:, the discoverer of 1954:and less often with 1869:Klinefelter syndrome 1424:polytene chromosomes 1072:Parascaris univalens 506:mitochondrial genome 5352:Genetics techniques 4980:Centromere position 4955:Polytene chromosome 4925:Circular chromosome 4298:2002PNAS...9916156W 4243:1996Sci...273..494S 4204:Histol. Histopathol 3950:1970Sci...168.1414C 3923:Volcanism in Hawaii 3762:1991PNAS...88.9051I 3243:Experientia (Basel) 3212:2005IchtR..52...94K 3130:1986Sci...231.1278C 3059:1970Sci...168.1364W 3012:10.1038/nature03479 3004:2005Natur.434..400C 2965:Gilbert S.F. 2006. 2937:2001JEZ...291..310W 2783:2000PNAS...97.9821G 2453:White M.J.D. 1973. 2048:Hans von Winiwarter 1846:and give rise to a 1790:virtual karyotyping 1786:Digital karyotyping 1781:Digital karyotyping 1221:, where one sex is 1079:n = 1; and an ant: 999:dosage compensation 843:, occurring in the 584:bands and sub-bands 424:staining is called 264:, is applied after 96:, and results in a 4753:Making a karyotype 3869:Systematic Biology 3255:10.1007/BF02153623 2473:. Academic Press. 2296:"Karyosystematics" 2135:Human chromosome 2 2103:hypotonic solution 2082: 2011:appearance of the 2003:, "general form") 1775: 1731: 1670:, less frequently 1657: 1547:Chromosome banding 1526:adaptive radiation 1516:All of the native 1357:Human chromosome 2 1116:Fundamental number 1097:shortnose sturgeon 813: 510: 490: 439:between the sexes, 245: 233: 147:and is designated 69: 58: 34:in the cells of a 5317: 5316: 5275: 5274: 5012:Centromere number 4929:Linear chromosome 4745:Media related to 4461:(10): 1063–1064. 4400:Molecular Biology 4188:978-3-8055-8019-9 4155:978-0-08-096156-9 4130:978-0-8248-2190-6 4103:978-1-4613-6877-9 3587:978-0-08-047052-8 3114:Myrmecia pilosula 3043:Muntiacus muntjak 2998:(7031): 400–404. 2116:by a solution of 2063:sex determination 1927:Angelman syndrome 1623:Silver staining: 1575:is obtained with 1447:subalpine meadows 1432:Hampton L. Carson 1374:Chilocorus stigma 1367:Some species are 1269:endoreduplication 1082:Myrmecia pilosula 1048:Muntiacus muntjak 1038:Muntiacus reevesi 1003:placental mammals 732: 731: 530:transcriptionally 306:amniotic centesis 290:white blood cells 272:by a solution of 98:photomicrographic 5359: 5142: 5141: 5106:Polyploidization 4934:Extra chromosome 4849:Genetic material 4812: 4805: 4798: 4789: 4788: 4744: 4728: 4722: 4716: 4710: 4704: 4703: 4693: 4669: 4663: 4662: 4648: 4642: 4641: 4621: 4615: 4614: 4602: 4596: 4595: 4583: 4577: 4576: 4556: 4550: 4543: 4537: 4536: 4527:(3–4): 193–197. 4516: 4510: 4505:Delaunay L. N. 4503: 4497: 4496: 4470: 4446: 4440: 4439: 4391: 4385: 4384: 4351:(8): 1973–1986. 4345:Nature Protocols 4336: 4330: 4329: 4319: 4309: 4292:(25): 16156–61. 4277: 4271: 4270: 4226: 4220: 4219: 4199: 4193: 4192: 4174: 4168: 4167: 4141: 4135: 4134: 4114: 4108: 4107: 4081: 4075: 4074: 4058: 4052: 4051: 4029: 4023: 4022: 4012: 3976: 3970: 3969: 3944:(3938): 1414–8. 3933: 3927: 3926: 3920: 3911: 3905: 3899: 3893: 3892: 3864: 3858: 3857: 3829: 3823: 3822: 3810: 3800: 3794: 3793: 3783: 3773: 3741: 3735: 3728: 3722: 3715: 3709: 3708: 3690: 3666: 3660: 3659: 3649: 3621: 3615: 3614: 3608: 3598: 3592: 3591: 3569: 3563: 3558: 3552: 3551: 3523: 3517: 3516: 3480: 3474: 3469: 3463: 3462: 3434: 3428: 3427: 3425: 3423: 3418:on 26 March 2012 3417: 3411:. Archived from 3394: 3381: 3375: 3374: 3372: 3354: 3341: 3335: 3334: 3312: 3306: 3305: 3303: 3273: 3267: 3266: 3238: 3232: 3231: 3191: 3185: 3184: 3164: 3158: 3157: 3109: 3103: 3096: 3087: 3086: 3053:(3937): 1364–6. 3038: 3032: 3031: 2987: 2981: 2976: 2970: 2963: 2957: 2956: 2945:10.1002/jez.1131 2920: 2914: 2913: 2877: 2871: 2870: 2834: 2828: 2821: 2815: 2814: 2804: 2794: 2762: 2756: 2755: 2749: 2741: 2731: 2707: 2701: 2700: 2694: 2686: 2654: 2648: 2647: 2641: 2633: 2623: 2613: 2589: 2583: 2582: 2545:Sax, L. (2002). 2542: 2536: 2535: 2529: 2521: 2511: 2491: 2485: 2484: 2464: 2458: 2451: 2442: 2439: 2430: 2429: 2419: 2409: 2385: 2376: 2375: 2359: 2349: 2343: 2336: 2327: 2324: 2318: 2316: 2306: 2300: 2299: 2292: 2286: 2285: 2273: 2263: 2257: 2251: 2242: 2241: 2235: 2225: 2219: 2218: 2200: 2194: 2193: 2191: 2189: 2173: 2158: 2058:, concluding an 2037:Michael JD White 2033:C. D. Darlington 2021:Grigory Levitsky 1977:Walther Flemming 1879:Edwards syndrome 1561:Types of banding 1461:, in the family 1439:Hawaiian Islands 1310:palaeopolyploidy 1280:nuclear membrane 1225:, and the other 1062: 941: 861:sister chromatid 621: 620: 208:karyosystematics 82:light microscopy 5367: 5366: 5362: 5361: 5360: 5358: 5357: 5356: 5322: 5321: 5318: 5313: 5271: 5178: 5140: 5110: 5099:Paleopolyploidy 5044: 5038: 4894: 4868:Heterochromatin 4831: 4825: 4816: 4783:Wayback Machine 4737: 4732: 4731: 4723: 4719: 4711: 4707: 4670: 4666: 4649: 4645: 4626:J. Exp. Zoology 4622: 4618: 4603: 4599: 4584: 4580: 4557: 4553: 4544: 4540: 4517: 4513: 4504: 4500: 4447: 4443: 4392: 4388: 4337: 4333: 4278: 4274: 4237:(5274): 494–7. 4227: 4223: 4200: 4196: 4189: 4175: 4171: 4156: 4142: 4138: 4131: 4115: 4111: 4104: 4082: 4078: 4059: 4055: 4048: 4030: 4026: 3977: 3973: 3934: 3930: 3918: 3912: 3908: 3900: 3896: 3865: 3861: 3840:(1–2): 68–169. 3830: 3826: 3819: 3801: 3797: 3742: 3738: 3729: 3725: 3716: 3712: 3667: 3663: 3640:(17): 2765–75. 3622: 3618: 3605:The chromosomes 3599: 3595: 3588: 3570: 3566: 3559: 3555: 3524: 3520: 3497:10.1038/nrg1711 3485:Nat. Rev. Genet 3481: 3477: 3470: 3466: 3435: 3431: 3421: 3419: 3415: 3392: 3382: 3378: 3352: 3342: 3338: 3313: 3309: 3274: 3270: 3239: 3235: 3192: 3188: 3165: 3161: 3110: 3106: 3098:Hsu T.C. 1979. 3097: 3090: 3039: 3035: 2988: 2984: 2977: 2973: 2964: 2960: 2921: 2917: 2878: 2874: 2835: 2831: 2822: 2818: 2763: 2759: 2743: 2742: 2708: 2704: 2688: 2687: 2655: 2651: 2635: 2634: 2590: 2586: 2543: 2539: 2523: 2522: 2492: 2488: 2481: 2467:K. Lew (2012). 2465: 2461: 2452: 2445: 2440: 2433: 2386: 2379: 2372: 2350: 2346: 2337: 2330: 2325: 2321: 2307: 2303: 2294: 2293: 2289: 2282: 2264: 2260: 2252: 2245: 2226: 2222: 2215: 2201: 2197: 2187: 2185: 2175: 2174: 2170: 2165: 2156: 2143: 2095:Using cells in 2029:Nikolai Vavilov 2025:Sergei Navashin 1987:in 1888. It is 1965: 1863:Turner syndrome 1856: 1804: 1798: 1783: 1723: 1648:probed for the 1638: 1604:Q-banding is a 1597:, so it stains 1563: 1558: 1549: 1420: 1365: 1334:Turner syndrome 1319: 1299:differentiation 1288: 1286: 1283: 1178: 1118: 1063: 1060: 1029:Kurt Benirschke 1021: 952:heterochromatin 948: 942: 939: 892: 870: 866: 852: 830: 820: 802: 604: 561:sex chromosomes 540:, and a higher 482: 480:Human karyogram 406:Giemsa staining 394:heterochromatin 318: 250: 217: 197:relationships, 172:sex chromosomes 130:sex chromosomes 24: 17: 12: 11: 5: 5365: 5355: 5354: 5349: 5344: 5339: 5334: 5315: 5314: 5312: 5311: 5306: 5301: 5296: 5295: 5294: 5283: 5281: 5277: 5276: 5273: 5272: 5270: 5269: 5264: 5259: 5254: 5249: 5244: 5239: 5234: 5229: 5224: 5219: 5214: 5209: 5204: 5199: 5194: 5188: 5186: 5180: 5179: 5177: 5176: 5171: 5166: 5161: 5156: 5150: 5148: 5139: 5138: 5133: 5118: 5116: 5112: 5111: 5109: 5108: 5103: 5102: 5101: 5096: 5091: 5086: 5076: 5075: 5074: 5069: 5059: 5054: 5048: 5046: 5040: 5039: 5037: 5036: 5035: 5034: 5029: 5024: 5019: 5009: 5008: 5007: 5002: 4997: 4992: 4990:Submetacentric 4987: 4977: 4972: 4967: 4962: 4957: 4952: 4947: 4942: 4937: 4931: 4922: 4917: 4916:or heterosome) 4910:Sex chromosome 4902: 4900: 4896: 4895: 4893: 4892: 4887: 4882: 4877: 4872: 4871: 4870: 4865: 4855: 4846: 4841: 4835: 4833: 4827: 4826: 4815: 4814: 4807: 4800: 4792: 4786: 4785: 4773: 4768: 4762: 4756: 4750: 4736: 4735:External links 4733: 4730: 4729: 4717: 4705: 4664: 4643: 4632:(3): 291–336. 4616: 4597: 4578: 4567:(4): 465–502. 4551: 4538: 4511: 4498: 4441: 4406:(4): 542–550. 4386: 4331: 4272: 4221: 4210:(1): 229–237. 4194: 4187: 4169: 4154: 4136: 4129: 4109: 4102: 4076: 4053: 4047:978-0849365478 4046: 4024: 3971: 3928: 3906: 3894: 3875:(2): 184–194. 3859: 3824: 3817: 3795: 3756:(20): 9051–5. 3736: 3723: 3717:Nagl W. 1978. 3710: 3681:(3): 297–306. 3661: 3616: 3593: 3586: 3564: 3553: 3518: 3491:(11): 836–46. 3475: 3464: 3451:10.1086/280872 3445:(750): 54–66. 3429: 3403:(1): 113–130. 3376: 3336: 3325:(2): 131–141. 3307: 3268: 3233: 3186: 3175:(3): 205–217. 3159: 3124:(4743): 1278. 3104: 3088: 3033: 2982: 2971: 2958: 2915: 2872: 2829: 2816: 2777:(18): 9821–3. 2757: 2702: 2649: 2584: 2557:(3): 174–178. 2537: 2486: 2479: 2459: 2443: 2431: 2377: 2370: 2344: 2328: 2319: 2301: 2287: 2280: 2258: 2243: 2220: 2213: 2195: 2167: 2166: 2164: 2161: 2160: 2159: 2150: 2142: 2139: 2127: 2126: 2123: 2120: 2106: 2099: 2097:tissue culture 1964: 1961: 1960: 1959: 1940: 1934: 1924: 1918: 1908: 1907: 1904: 1898: 1895:Patau syndrome 1892: 1886: 1876: 1866: 1855: 1852: 1848:genetic mosaic 1824:nondisjunction 1812:translocations 1800:Main article: 1797: 1794: 1782: 1779: 1755:interferometer 1722: 1719: 1637: 1634: 1633: 1632: 1625:Silver nitrate 1621: 1614: 1602: 1591: 1588: 1562: 1559: 1557: 1554: 1548: 1545: 1497:moving over a 1419: 1416: 1399:Thais lapillus 1364: 1361: 1318: 1315: 1314: 1313: 1306: 1241:differentiated 1237:Endopolyploidy 1234: 1219:Haplo-diploidy 1208: 1207: 1177: 1174: 1117: 1114: 1058: 1020: 1017: 1016: 1015: 991:X-inactivation 988: 967:Theodor Boveri 963: 947: 944: 937: 912:macromolecules 891: 888: 868: 864: 850: 828: 818: 801: 798: 797: 796: 785: 758:sex chromosome 754: 738:versus inside 730: 729: 722: 719: 713: 712: 709: 706: 700: 699: 696: 693: 687: 686: 680: 677: 671: 670: 667: 664: 658: 657: 654: 651: 645: 644: 641: 638: 632: 631: 628: 625: 603: 600: 538:non-coding DNA 481: 478: 477: 476: 469: 462: 455: 440: 430: 429: 366: 359: 355: 352: 349:translocations 341: 317: 314: 294:tissue culture 249: 246: 216: 213: 162:So, in normal 145:somatic number 15: 9: 6: 4: 3: 2: 5364: 5353: 5350: 5348: 5345: 5343: 5340: 5338: 5335: 5333: 5330: 5329: 5327: 5320: 5310: 5307: 5305: 5302: 5300: 5297: 5293: 5290: 5289: 5288: 5285: 5284: 5282: 5278: 5268: 5265: 5263: 5260: 5258: 5255: 5253: 5250: 5248: 5245: 5243: 5240: 5238: 5235: 5233: 5230: 5228: 5225: 5223: 5220: 5218: 5215: 5213: 5210: 5208: 5205: 5203: 5200: 5198: 5195: 5193: 5190: 5189: 5187: 5185: 5181: 5175: 5172: 5170: 5167: 5165: 5162: 5160: 5157: 5155: 5152: 5151: 5149: 5147: 5143: 5137: 5134: 5131: 5127: 5123: 5120: 5119: 5117: 5113: 5107: 5104: 5100: 5097: 5095: 5092: 5090: 5087: 5085: 5082: 5081: 5080: 5077: 5073: 5070: 5068: 5065: 5064: 5063: 5060: 5058: 5055: 5053: 5050: 5049: 5047: 5045:and evolution 5041: 5033: 5030: 5028: 5025: 5023: 5020: 5018: 5015: 5014: 5013: 5010: 5006: 5003: 5001: 4998: 4996: 4993: 4991: 4988: 4986: 4983: 4982: 4981: 4978: 4976: 4973: 4971: 4970:Isochromosome 4968: 4966: 4963: 4961: 4958: 4956: 4953: 4951: 4948: 4946: 4943: 4941: 4938: 4935: 4932: 4930: 4926: 4923: 4921: 4918: 4915: 4911: 4907: 4904: 4903: 4901: 4897: 4891: 4888: 4886: 4883: 4881: 4878: 4876: 4873: 4869: 4866: 4864: 4861: 4860: 4859: 4856: 4854: 4850: 4847: 4845: 4842: 4840: 4837: 4836: 4834: 4828: 4824: 4820: 4813: 4808: 4806: 4801: 4799: 4794: 4793: 4790: 4784: 4780: 4777: 4774: 4772: 4769: 4766: 4763: 4760: 4757: 4754: 4751: 4748: 4743: 4739: 4738: 4726: 4721: 4714: 4709: 4701: 4697: 4692: 4687: 4683: 4679: 4675: 4668: 4660: 4659: 4654: 4647: 4639: 4635: 4631: 4627: 4620: 4612: 4608: 4601: 4593: 4589: 4582: 4574: 4570: 4566: 4562: 4561:Bull Hist Med 4555: 4548: 4542: 4534: 4530: 4526: 4522: 4515: 4508: 4502: 4494: 4490: 4486: 4482: 4478: 4474: 4469: 4464: 4460: 4456: 4452: 4445: 4437: 4433: 4429: 4425: 4421: 4417: 4413: 4409: 4405: 4401: 4397: 4390: 4382: 4378: 4374: 4370: 4366: 4362: 4358: 4354: 4350: 4346: 4342: 4335: 4327: 4323: 4318: 4313: 4308: 4303: 4299: 4295: 4291: 4287: 4283: 4276: 4268: 4264: 4260: 4256: 4252: 4248: 4244: 4240: 4236: 4232: 4225: 4217: 4213: 4209: 4205: 4198: 4190: 4184: 4180: 4173: 4165: 4161: 4157: 4151: 4147: 4140: 4132: 4126: 4122: 4121: 4113: 4105: 4099: 4095: 4091: 4087: 4080: 4072: 4068: 4064: 4057: 4049: 4043: 4039: 4035: 4028: 4020: 4016: 4011: 4006: 4002: 3998: 3995:(3): 465–82. 3994: 3990: 3986: 3984: 3975: 3967: 3963: 3959: 3955: 3951: 3947: 3943: 3939: 3932: 3924: 3917: 3910: 3904:, p. 169 3903: 3898: 3890: 3886: 3882: 3878: 3874: 3870: 3863: 3855: 3851: 3847: 3843: 3839: 3835: 3828: 3820: 3818:9780387076683 3814: 3809: 3808: 3799: 3791: 3787: 3782: 3777: 3772: 3767: 3763: 3759: 3755: 3751: 3747: 3740: 3733: 3727: 3720: 3714: 3706: 3702: 3698: 3694: 3689: 3684: 3680: 3676: 3672: 3665: 3657: 3653: 3648: 3643: 3639: 3635: 3631: 3629: 3620: 3612: 3607: 3606: 3597: 3589: 3583: 3579: 3575: 3568: 3562: 3561:Stebbins 1971 3557: 3549: 3545: 3541: 3537: 3534:(2): 135–41. 3533: 3529: 3522: 3514: 3510: 3506: 3502: 3498: 3494: 3490: 3486: 3479: 3473: 3472:Stebbins 1950 3468: 3460: 3456: 3452: 3448: 3444: 3440: 3433: 3414: 3410: 3406: 3402: 3398: 3391: 3389: 3380: 3371: 3366: 3362: 3358: 3351: 3349: 3340: 3332: 3328: 3324: 3320: 3319: 3311: 3302: 3297: 3293: 3289: 3288: 3283: 3281: 3272: 3264: 3260: 3256: 3252: 3248: 3244: 3237: 3229: 3225: 3221: 3217: 3213: 3209: 3205: 3201: 3197: 3190: 3182: 3178: 3174: 3170: 3163: 3155: 3151: 3147: 3143: 3139: 3135: 3131: 3127: 3123: 3119: 3115: 3108: 3101: 3095: 3093: 3084: 3080: 3076: 3072: 3068: 3064: 3060: 3056: 3052: 3048: 3044: 3037: 3029: 3025: 3021: 3017: 3013: 3009: 3005: 3001: 2997: 2993: 2986: 2980: 2975: 2968: 2962: 2954: 2950: 2946: 2942: 2938: 2934: 2930: 2926: 2919: 2911: 2907: 2903: 2899: 2895: 2891: 2887: 2883: 2876: 2868: 2864: 2860: 2856: 2852: 2848: 2845:(3): 242–50. 2844: 2840: 2833: 2826: 2820: 2812: 2808: 2803: 2798: 2793: 2788: 2784: 2780: 2776: 2772: 2768: 2761: 2753: 2747: 2739: 2735: 2730: 2725: 2721: 2717: 2713: 2706: 2698: 2692: 2684: 2680: 2676: 2672: 2669:(3): 1360–5. 2668: 2664: 2660: 2653: 2645: 2639: 2631: 2627: 2622: 2617: 2612: 2607: 2603: 2599: 2595: 2588: 2580: 2576: 2572: 2568: 2564: 2560: 2556: 2552: 2548: 2541: 2533: 2527: 2519: 2515: 2510: 2505: 2501: 2497: 2490: 2482: 2480:9780123813749 2476: 2472: 2471: 2463: 2456: 2450: 2448: 2438: 2436: 2427: 2423: 2418: 2413: 2408: 2403: 2399: 2395: 2391: 2384: 2382: 2373: 2371:9780713122879 2367: 2363: 2358: 2357: 2348: 2341: 2335: 2333: 2323: 2314: 2313: 2305: 2297: 2291: 2283: 2281:9780231017336 2277: 2272: 2271: 2262: 2255: 2250: 2248: 2239: 2234: 2233: 2224: 2216: 2214:0-87893-403-0 2210: 2206: 2199: 2184: 2183: 2178: 2172: 2168: 2154: 2153:Genome screen 2151: 2148: 2145: 2144: 2138: 2136: 2132: 2124: 2121: 2119: 2115: 2111: 2107: 2104: 2100: 2098: 2094: 2093: 2092: 2090: 2086: 2078: 2074: 2072: 2068: 2064: 2061: 2057: 2053: 2052:spermatogonia 2049: 2045: 2040: 2038: 2034: 2030: 2026: 2022: 2018: 2014: 2010: 2004: 2002: 1998: 1994: 1993:Ancient Greek 1990: 1986: 1982: 1978: 1974: 1970: 1957: 1953: 1949: 1945: 1941: 1938: 1935: 1932: 1928: 1925: 1922: 1919: 1916: 1913: 1912: 1911: 1905: 1902: 1899: 1896: 1893: 1890: 1889:Down syndrome 1887: 1884: 1881:is caused by 1880: 1877: 1874: 1870: 1867: 1864: 1861: 1860: 1859: 1851: 1849: 1845: 1841: 1837: 1833: 1829: 1825: 1821: 1817: 1813: 1809: 1803: 1793: 1791: 1787: 1778: 1771: 1767: 1765: 1760: 1756: 1752: 1748: 1747:Fluorescently 1744: 1740: 1736: 1727: 1718: 1716: 1712: 1708: 1704: 1700: 1696: 1691: 1689: 1685: 1681: 1677: 1673: 1669: 1666: 1662: 1655: 1651: 1647: 1642: 1630: 1626: 1622: 1619: 1615: 1611: 1607: 1603: 1600: 1596: 1592: 1589: 1586: 1582: 1578: 1574: 1571: 1570: 1569: 1567: 1553: 1544: 1541: 1539: 1535: 1531: 1527: 1523: 1519: 1514: 1512: 1508: 1504: 1500: 1496: 1495:Pacific Plate 1492: 1488: 1484: 1480: 1474: 1472: 1466: 1464: 1463:Drosophilidae 1460: 1459: 1454: 1453: 1448: 1444: 1440: 1435: 1433: 1429: 1425: 1418:Species trees 1415: 1413: 1409: 1405: 1401: 1400: 1396: 1392: 1391: 1390:Sorex araneus 1386: 1385: 1381:of the genus 1380: 1376: 1375: 1370: 1360: 1358: 1354: 1350: 1349: 1344: 1343: 1337: 1335: 1331: 1330:Down syndrome 1327: 1323: 1311: 1307: 1305:in many ways. 1304: 1303:morphogenesis 1300: 1296: 1293:kingdom from 1292: 1281: 1278: 1274: 1273:reduplication 1270: 1266: 1262: 1258: 1254: 1250: 1246: 1242: 1238: 1235: 1232: 1228: 1224: 1220: 1217: 1216: 1215: 1213: 1205: 1201: 1197: 1193: 1189: 1186: 1185: 1184: 1182: 1173: 1171: 1167: 1163: 1159: 1155: 1151: 1147: 1143: 1139: 1135: 1131: 1127: 1123: 1113: 1111: 1107: 1106:B chromosomes 1103: 1102: 1098: 1094: 1093: 1088: 1084: 1083: 1078: 1074: 1073: 1069: 1057: 1052: 1050: 1049: 1044: 1040: 1039: 1034: 1033:Doris Wurster 1030: 1026: 1013: 1008: 1004: 1000: 996: 992: 989: 986: 982: 981: 976: 972: 968: 964: 961: 960:sciarid flies 957: 956: 955: 953: 940:Maynard Smith 936: 932: 927: 922: 919: 917: 913: 909: 905: 901: 900:transcription 897: 894:Although the 887: 885: 881: 876: 874: 862: 858: 854: 846: 842: 841:DNA synthesis 838: 834: 831:phase of the 826: 822: 811: 806: 794: 790: 786: 783: 779: 775: 771: 770:X chromosomes 767: 763: 759: 755: 752: 749: 745: 744: 743: 741: 737: 727: 723: 720: 718: 715: 714: 710: 707: 705: 702: 701: 697: 694: 692: 689: 688: 685: 681: 678: 676: 673: 672: 668: 665: 663: 660: 659: 655: 652: 650: 647: 646: 642: 639: 637: 634: 633: 629: 626: 623: 622: 619: 617: 613: 609: 599: 597: 593: 589: 585: 581: 576: 574: 570: 566: 562: 558: 554: 550: 545: 543: 539: 535: 531: 527: 523: 519: 515: 507: 503: 499: 494: 486: 474: 470: 467: 463: 460: 456: 453: 449: 445: 441: 438: 437: 436: 433: 427: 423: 419: 415: 411: 407: 403: 399: 395: 391: 387: 383: 380:pairs versus 379: 375: 371: 367: 364: 360: 356: 353: 350: 346: 342: 339: 335: 334: 329: 328: 323: 322: 321: 313: 311: 307: 303: 299: 295: 291: 287: 283: 279: 275: 271: 270:cell division 267: 263: 259: 255: 242: 237: 230: 226: 221: 212: 210: 209: 204: 200: 196: 192: 188: 183: 181: 177: 173: 169: 165: 160: 158: 154: 150: 146: 142: 137: 135: 131: 127: 123: 119: 114: 112: 107: 103: 99: 95: 91: 87: 83: 78: 74: 66: 62: 55: 51: 47: 43: 41: 37: 33: 29: 22: 5342:Cytogenetics 5332:Cell biology 5319: 5011: 4979: 4838: 4819:Cytogenetics 4727:(video) 2007 4720: 4708: 4684:(1–2): 1–6. 4681: 4677: 4667: 4658:The Guardian 4656: 4646: 4629: 4625: 4619: 4610: 4606: 4600: 4594:(93): 147–9. 4591: 4587: 4581: 4564: 4560: 4554: 4546: 4541: 4524: 4520: 4514: 4506: 4501: 4458: 4454: 4444: 4403: 4399: 4389: 4348: 4344: 4334: 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