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Chromosome
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Diagram of a replicated and condensed metaphase eukaryotic chromosome. (1) Chromatid – one of the two identical parts of the chromosome after S phase. (2) Centromere – the point where the two chromatids touch. (3) Short (p) arm. (4) Long (q) arm.
A chromosome (from ancient Greek: χρωμόσωμα, chromosoma, chroma means colour, soma means body) is a DNA molecule with part or all of the genetic material (genome) of an organism. Most eukaryotic chromosomes include packaging proteins which, aided by chaperone proteins, bind to and condense the DNA molecule to prevent it from becoming an unmanageable tangle.[1][2]
Chromosomes are normally visible under a light microscope only when the cell is undergoing the metaphase of cell division (where all chromosomes are aligned in the center of the cell in their condensed form).[3] Before this happens, every chromosome is copied once (S phase), and the copy is joined to the original by a centromere, resulting either in an X-shaped structure (pictured to the right) if the centromere is located in the middle of the chromosome or a two-arm structure if the centromere is located near one of the ends. The original chromosome and the copy are now called sister chromatids. During metaphase the X-shape structure is called a metaphase chromosome. In this highly condensed form chromosomes are easiest to distinguish and study.[4] In animal cells, chromosomes reach their highest compaction level in anaphase during segregation.[5]
Chromosomal recombination during meiosis and subsequent sexual reproduction play a significant role in genetic diversity. If these structures are manipulated incorrectly, through processes known as chromosomal instability and translocation, the cell may undergo mitotic catastrophe and die. Mutations in the cell can allow it to inappropriately evade apoptosis and lead to the progression of cancer.
Some use the term chromosome in a wider sense, to refer to the individualized portions of chromatin in cells, either visible or not under light microscopy. Others use the concept in a narrower sense, to refer to the individualized portions of chromatin during cell division, visible under light microscopy due to high condensation.
Contents
[hide]
1 Etymology
2 History of discovery
3 Prokaryotes
3.1 Structure in sequences
3.2 DNA packaging
4 Eukaryotes
4.1 Interphase chromatin
4.2 Metaphase chromatin and division
4.3 Human chromosomes
5 Number in various organisms
5.1 In eukaryotes
5.2 In prokaryotes
6 Karyotype
6.1 Historical note
7 Aberrations
7.1 Sperm aneuploidy
8 See also
9 Notes and references
10 External links
Etymology[edit]
The word chromosome (/ˈkroʊməˌsoʊm, -ˌzoʊm/[6][7]) comes from the Greek χρῶμα (chroma, "colour") and σῶμα (soma, "body"), describing their strong staining by particular dyes.[8] The term was coined by von Waldeyer-Hartz,[9] referring to the term chromatin, which was introduced by Walther Flemming.
Emilio Battaglia (1917-2011)[10][11] points out that over time many of the most familiar caryological terms have become inadequate or illogical or, in some cases, etymologically incorrect so that they should be replaced by more adequate alternatives suggested by the present scientific progress. The author has been particularly disappointed by the illogicality of the present chromosomal (chromatin-chromosome) terminology based on, or inferred by, two terms, Chromatin (Flemming 1880) and Chromosom (Waldeyer 1888), both inappropriately ascribed to a basically non coloured state.[12]
History of discovery[edit]
Walter Sutton (left) and Theodor Boveri (right) independently developed the chromosome theory of inheritance in 1902.
Schleiden,[4] Virchow and Bütschli were among the first scientists who recognized the structures now familiar as chromosomes.[13]
In a series of experiments beginning in the mid-1880s, Theodor Boveri gave the definitive demonstration that chromosomes are the vectors of heredity. His two principles were the continuity of chromosomes and the individuality of chromosomes.[citation needed][further explanation needed] It is the second of these principles that was so original.[citation needed] Wilhelm Roux suggested that each chromosome carries a different genetic load. Boveri was able to test and confirm this hypothesis. Aided by the rediscovery at the start of the 1900s of Gregor Mendel's earlier work, Boveri was able to point out the connection between the rules of inheritance and the behaviour of the chromosomes. Boveri influenced two generations of American cytologists: Edmund Beecher Wilson, Nettie Stevens, Walter Sutton and Theophilus Painter were all influenced by Boveri (Wilson, Stevens, and Painter actually worked with him).[14]
In his famous textbook The Cell in Development and Heredity, Wilson linked together the independent work of Boveri and Sutton (both around 1902) by naming the chromosome theory of inheritance the Boveri–Sutton chromosome theory (the names are sometimes reversed).[15] Ernst Mayr remarks that the theory was hotly contested by some famous geneticists: William Bateson, Wilhelm Johannsen, Richard Goldschmidt and T.H. Morgan, all of a rather dogmatic turn of mind. Eventually, complete proof came from chromosome maps in Morgan's own lab.[16]
The number of human chromosomes was published in 1923 by Theophilus Painter. By inspection through the microscope, he counted 24 pairs, which would mean 48 chromosomes. His error was copied by others and it was not until 1956 that the true number, 46, was determined by Indonesia-born cytogeneticist Joe Hin Tjio.[17]
Prokaryotes[edit]
The prokaryotes – bacteria and archaea – typically have a single circular chromosome, but many variations exist.[18] The chromosomes of most bacteria, which some authors prefer to call genophores, can range in size from only 130,000 base pairs in the endosymbiotic bacteria Candidatus Hodgkinia cicadicola[19] and Candidatus Tremblaya princeps,[20] to more than 14,000,000 base pairs in the soil-dwelling bacterium Sorangium cellulosum.[21] Spirochaetes of the genus Borrelia are a notable exception to this arrangement, with bacteria such as Borrelia burgdorferi, the cause of Lyme disease, containing a single linear chromosome.[22]
Structure in sequences[edit]
Prokaryotic chromosomes have less sequence-based structure than eukaryotes. Bacteria typically have a one-point (the origin of replication) from which replication starts, whereas some archaea contain multiple replication origins.[23] The genes in prokaryotes are often organized in operons, and do not usually contain introns, unlike eukaryotes.
DNA packaging[edit]
Prokaryotes do not possess nuclei. Instead, their DNA is organized into a structure called the nucleoid.[24][25] The nucleoid is a distinct structure and occupies a defined region of the bacterial cell. This structure is, however, dynamic and is maintained and remodeled by the actions of a range of histone-like proteins, which associate with the bacterial chromosome.[26] In archaea, the DNA in chromosomes is even more organized, with the DNA packaged within structures similar to eukaryotic nucleosomes.[27][28]
Certain bacteria also contain plasmids or other extrachromosomal DNA. These are circular structures in the cytoplasm that contain cellular DNA and play a role in horizontal gene transfer.[4] In prokaryotes (see nucleoids) and viruses,[29] the DNA is often densely packed and organized; in the case of archaea, by homology to eukaryotic histones, and in the case of bacteria, by histone-like proteins.
Bacterial chromosomes tend to be tethered to the plasma membrane of the bacteria. In molecular biology application, this allows for its isolation from plasmid DNA by centrifugation of lysed bacteria and pelleting of the membranes (and the attached DNA).
Prokaryotic chromosomes and plasmids are, like eukaryotic DNA, generally supercoiled. The DNA must first be released into its relaxed state for access for transcription, regulation, and replication.
Eukaryotes[edit]
Organization of DNA in a eukaryotic cell.
See also: Eukaryotic chromosome fine structure
Chromosomes in eukaryotes are composed of chromatin fiber. Chromatin fiber is made of nucleosomes (histone octamers with part of a DNA strand attached to and wrapped around it). Chromatin fibers are packaged by proteins into a condensed structure called chromatin. Chromatin contains the vast majority of DNA and a small amount inherited maternally, can be found in the mitochondria. Chromatin is present in most cells, with a few exceptions, for example, red blood cells.
Chromatin allows the very long DNA molecules to fit into the cell nucleus. During cell division chromatin condenses further to form microscopically visible chromosomes. The structure of chromosomes varies through the cell cycle. During cellular division chromosomes are replicated, divided, and passed successfully to their daughter cells so as to ensure the genetic diversity and survival of their progeny. Chromosomes may exist as either duplicated or unduplicated. Unduplicated chromosomes are single double helixes, whereas duplicated chromosomes contain two identical copies (called chromatids or sister chromatids) joined by a centromere.
The major structures in DNA compaction: DNA, the nucleosome, the 10 nm "beads-on-a-string" fibre, the 30 nm fibre and the metaphase chromosome.
Eukaryotes (cells with nuclei such as those found in plants, fungi, and animals) possess multiple large linear chromosomes contained in the cell's nucleus. Each chromosome has one centromere, with one or two arms projecting from the centromere, although, under most circumstances, these arms are not visible as such. In addition, most eukaryotes have a small circular mitochondrial genome, and some eukaryotes may have additional small circular or linear cytoplasmic chromosomes.
In the nuclear chromosomes of eukaryotes, the uncondensed DNA exists in a semi-ordered structure, where it is wrapped around histones (structural proteins), forming a composite material called chromatin.
Interphase chromatin[edit]
During interphase (the period of the cell cycle where the cell is not dividing), two types of chromatin can be distinguished:
Euchromatin, which consists of DNA that is active, e.g., being expressed as protein.
Heterochromatin, which consists of mostly inactive DNA. It seems to serve structural purposes during the chromosomal stages. Heterochromatin can be further distinguished into two types:
Constitutive heterochromatin, which is never expressed. It is located around the centromere and usually contains repetitive sequences.
Facultative heterochromatin, which is sometimes expressed.
Structure of Eukaryotic chromosome
Each chromosome is made up of two chromatids(chromosomal arms) which are joined to each other at a small constricted region called the centromere.(Primary constriction). These sister chromatids are conjoined twins the result of DNA replication.
The centromere helps the chromatids attach to the spindle fibres during cell division, it is also concerned with the anaphase movement of the chromosomes, by which the spindle fibers pull the chromatids to the two opposite poles by their contraction during anaphase.
Besides the primary constriction, in certain chromosomes there is a secondary constriction as well. Because a small portion is pinched off from the chromosomal body; this portion is called a 'satellite' and the chromosome is called an SAT chromosome.
The two chromatids are made up of very thin chromatin fibres which are made up of 40% DNA and 60% histone proteins
Each chromatin fibre consists of one DNA helix coiled around eight histone molecules like a loop; such a complex is called nucleosome and resembles a bead on a string. These nucleosomes pack tighter, during condensation required to get to metaphase.
The primary constriction cannot take up most stains, so during cell division this region is a gap in staining.
Within the primary constriction there is a clear zone called Centromere.
The centromere with the DNA and histone proteins bound to them form a disc shaped structure called kinetochore.
the chromonemata is a word that means a chromatid in the early stage of condensation.
Metaphase chromatin and division[edit]
See also: mitosis and meiosis
Human chromosomes during metaphase
In the early stages of mitosis or meiosis (cell division), the chromatin double helix become more and more condensed. They cease to function as accessible genetic material (transcription stops) and become a compact transportable form. This compact form makes the individual chromosomes visible, and they form the classic four arm structure, a pair of sister chromatids attached to each other at the centromere. The shorter arms are called p arms (from the French petit, small) and the longer arms are called q arms (q follows p in the Latin alphabet; q-g "grande"; alternatively it is sometimes said q is short for queue meaning tail in French[30]). This is the only natural context in which individual chromosomes are visible with an optical microscope.
Mitotic metaphase chromosomes are best described by a linearly organized longitudinally compressed array of consecutive chromatin loops.[31]
During mitosis, microtubules grow from centrosomes located at opposite ends of the cell and also attach to the centromere at specialized structures called kinetochores, one of which is present on each sister chromatid. A special DNA base sequence in the region of the kinetochores provides, along with special proteins, longer-lasting attachment in this region. The microtubules then pull the chromatids apart toward the centrosomes, so that each daughter cell inherits one set of chromatids. Once the cells have divided, the chromatids are uncoiled and DNA can again be transcribed. In spite of their appearance, chromosomes are structurally highly condensed, which enables these giant DNA structures to be contained within a cell nucleus.
Human chromosomes[edit]
Chromosomes in humans can be divided into two types: autosomes (body chromosome(s)) and allosome (sex chromosome(s)). Certain genetic traits are linked to a person's sex and are passed on through the sex chromosomes. The autosomes contain the rest of the genetic hereditary information. All act in the same way during cell division. Human cells have 23 pairs of chromosomes (22 pairs of autosomes and one pair of sex chromosomes), giving a total of 46 per cell. In addition to these, human cells have many hundreds of copies of the mitochondrial genome. Sequencing of the human genome has provided a great deal of information about each of the chromosomes. Below is a table compiling statistics for the chromosomes, based on the Sanger Institute's human genome information in the Vertebrate Genome Annotation (VEGA) database.[32] Number of genes is an estimate, as it is in part based on gene predictions. Total chromosome length is an estimate as well, based on the estimated size of unsequenced heterochromatin regions.
Estimated number of genes and base pairs (in mega base pairs) on each human chromosome
Chromosome
Genes[33]
Total base pairs
% of bases
Sequenced base pairs[34]
1 2000 247,199,719 8.0 224,999,719
2 1300 242,751,149 7.9 237,712,649
3 1000 199,446,827 6.5 194,704,827
4 1000 191,263,063 6.2 187,297,063
5 900 180,837,866 5.9 177,702,766
6 1000 170,896,993 5.5 167,273,993
7 900 158,821,424 5.2 154,952,424
8 700 146,274,826 4.7 142,612,826
9 800 140,442,298 4.6 120,312,298
10 700 135,374,737 4.4 131,624,737
11 1300 134,452,384 4.4 131,130,853
12 1100 132,289,534 4.3 130,303,534
13 300 114,127,980 3.7 95,559,980
14 800 106,360,585 3.5 88,290,585
15 600 100,338,915 3.3 81,341,915
16 800 88,822,254 2.9 78,884,754
17 1200 78,654,742 2.6 77,800,220
18 200 76,117,153 2.5 74,656,155
19 1500 63,806,651 2.1 55,785,651
20 500 62,435,965 2.0 59,505,254
21 200 46,944,323 1.5 34,171,998
22 500 49,528,953 1.6 34,893,953
X (sex chromosome) 800 154,913,754 5.0 151,058,754
Y (sex chromosome) 50 57,741,652 1.9 25,121,652
Total
21,000
3,079,843,747
100.0
2,857,698,560
Number in various organisms[edit]
Main article: List of organisms by chromosome count
In eukaryotes[edit]
These tables give the total number of chromosomes (including sex chromosomes) in a cell nucleus. For example, most eukaryotes are diploid, like humans who have 22 different types of autosomes, each present as two homologous pairs, and two sex chromosomes. This gives 46 chromosomes in total. Other organisms have more than two copies of their chromosome types, such as bread wheat, which is hexaploid and has six copies of seven different chromosome types – 42 chromosomes in total.
Chromosome numbers in some plants
Plant Species
#
Arabidopsis thaliana (diploid)[35] 10
Rye (diploid)[36] 14
Einkorn wheat (diploid)[37] 14
Maize (diploid or palaeotetraploid)[38] 20
Durum wheat (tetraploid)[37] 28
Bread wheat (hexaploid)[37] 42
Cultivated tobacco (tetraploid)[39] 48
Adder's tongue fern (polyploid)[40] approx. 1,200
Chromosome numbers (2n) in some animals
Species
#
Indian muntjac 7
Common fruit fly 8
Pill millipede (Arthrosphaera fumosa)[41] 30
Earthworm (Octodrilus complanatus)[42] 36
Tibetan fox 36
Domestic cat[43] 38
Domestic pig 38
Laboratory mouse[44][45] 40
Laboratory rat[45] 42
Rabbit (Oryctolagus cuniculus)[46] 44
Syrian hamster[44] 44
Guppy (poecilia reticulata)[47] 46
Human[48] 46
Hares[49][50] 48
Gorillas, chimpanzees[48] 48
Domestic sheep 54
Garden snail[51] 54
Silkworm[52] 56
Elephants[53] 56
Cow 60
Donkey 62
Guinea pig[54] 64
Horse 64
Dog[55] 78
Hedgehog 90
Goldfish[56] 100-104
Kingfisher[57] 132
Chromosome numbers in other organisms
Species
Large
Chromosomes
Intermediate
Chromosomes
Microchromosomes
Trypanosoma brucei 11 6 ~100
Domestic pigeon
(Columba livia domestics)[58] 18 - 59-63
Chicken[59] 8 2 sex chromosomes 60
Normal members of a particular eukaryotic species all have the same number of nuclear chromosomes (see the table). Other eukaryotic chromosomes, i.e., mitochondrial and plasmid-like small chromosomes, are much more variable in number, and there may be thousands of copies per cell.
The 23 human chromosome territories during prometaphase in fibroblast cells.
Asexually reproducing species have one set of chromosomes that are the same in all body cells. However, asexual species can be either haploid or diploid.
Sexually reproducing species have somatic cells (body cells), which are diploid [2n] having two sets of chromosomes (23 pairs in humans with one set of 23 chromosomes from each parent), one set from the mother and one from the father. Gametes, reproductive cells, are haploid [n]: They have one set of chromosomes. Gametes are produced by meiosis of a diploid germ line cell. During meiosis, the matching chromosomes of father and mother can exchange small parts of themselves (crossover), and thus create new chromosomes that are not inherited solely from either parent. When a male and a female gamete merge (fertilization), a new diploid organism is formed.
Some animal and plant species are polyploid [Xn]: They have more than two sets of homologous chromosomes. Plants important in agriculture such as tobacco or wheat are often polyploid, compared to their ancestral species. Wheat has a haploid number of seven chromosomes, still seen in some cultivars as well as the wild progenitors. The more-common pasta and bread wheat types are polyploid, having 28 (tetraploid) and 42 (hexaploid) chromosomes, compared to the 14 (diploid) chromosomes in the wild wheat.[60]
In prokaryotes[edit]
Prokaryote species generally have one copy of each major chromosome, but most cells can easily survive with multiple copies.[61] For example, Buchnera, a symbiont of aphids has multiple copies of its chromosome, ranging from 10–400 copies per cell.[62] However, in some large bacteria, such as Epulopiscium fishelsoni up to 100,000 copies of the chromosome can be present.[63] Plasmids and plasmid-like small chromosomes are, as in eukaryotes, highly variable in copy number. The number of plasmids in the cell is almost entirely determined by the rate of division of the plasmid – fast division causes high copy number.
Karyotype[edit]
Main article: Karyotype
Karyogram of a human male
In general, the karyotype is the characteristic chromosome complement of a eukaryote species.[64] The preparation and study of karyotypes is part of cytogenetics.
Although the replication and transcription of DNA is highly standardized in eukaryotes, the same cannot be said for their karyotypes, which are often highly variable. There may be variation between species in chromosome number and in detailed organization. In some cases, there is significant variation within species. Often there is:
1. variation between the two sexes
2. variation between the germ-line and soma (between gametes and the rest of the body)
3. variation between members of a population, due to balanced genetic polymorphism
4. geographical variation between races
5. mosaics or otherwise abnormal individuals.
Also, variation in karyotype may occur during development from the fertilized egg.
The technique of determining the karyotype is usually called karyotyping. Cells can be locked part-way through division (in metaphase) in vitro (in a reaction vial) with colchicine. These cells are then stained, photographed, and arranged into a karyogram, with the set of chromosomes arranged, autosomes in order of length, and sex chromosomes (here X/Y) at the end.
Like many sexually reproducing species, humans have special gonosomes (sex chromosomes, in contrast to autosomes). These are XX in females and XY in males.
Historical note[edit]
See also: Argument from authority § Use in science
Investigation into the human karyotype took many years to settle the most basic question: How many chromosomes does a normal diploid human cell contain? In 1912, Hans von Winiwarter reported 47 chromosomes in spermatogonia and 48 in oogonia, concluding an XX/XO sex determination mechanism.[65] Painter in 1922 was not certain whether the diploid number of man is 46 or 48, at first favouring 46.[66] He revised his opinion later from 46 to 48, and he correctly insisted on humans having an XX/XY system.[67]
New techniques were needed to definitively solve the problem:
Using cells in culture
Arresting mitosis in metaphase by a solution of colchicine
Pretreating cells in a hypotonic solution 0.075 M KCl, which swells them and spreads the chromosomes
Squashing the preparation on the slide forcing the chromosomes into a single plane
Cutting up a photomicrograph and arranging the result into an indisputable karyogram.
It took until 1954 before the human diploid number was confirmed as 46.[68][69] Considering the techniques of Winiwarter and Painter, their results were quite remarkable.[70] Chimpanzees, the closest living relatives to modern humans, have 48 chromosomes as do the other great apes: in humans two chromosomes fused to form chromosome 2.
Aberrations[edit]
In Down syndrome, there are three copies of chromosome 21
Chromosomal aberrations are disruptions in the normal chromosomal content of a cell and are a major cause of genetic conditions in humans, such as Down syndrome, although most aberrations have little to no effect. Some chromosome abnormalities do not cause disease in carriers, such as translocations, or chromosomal inversions, although they may lead to a higher chance of bearing a child with a chromosome disorder. Abnormal numbers of chromosomes or chromosome sets, called aneuploidy, may be lethal or may give rise to genetic disorders.[71] Genetic counseling is offered for families that may carry a chromosome rearrangement.
The gain or loss of DNA from chromosomes can lead to a variety of genetic disorders. Human examples include:
Cri du chat, which is caused by the deletion of part of the short arm of chromosome 5. "Cri du chat" means "cry of the cat" in French; the condition was so-named because affected babies make high-pitched cries that sound like those of a cat. Affected individuals have wide-set eyes, a small head and jaw, moderate to severe mental health problems, and are very short.
Down syndrome, the most common trisomy, usually caused by an extra copy of chromosome 21 (trisomy 21). Characteristics include decreased muscle tone, stockier build, asymmetrical skull, slanting eyes and mild to moderate developmental disability.[72]
Edwards syndrome, or trisomy-18, the second most common trisomy.[73] Symptoms include motor retardation, developmental disability and numerous congenital anomalies causing serious health problems. Ninety percent of those affected die in infancy. They have characteristic clenched hands and overlapping fingers.
Isodicentric 15, also called idic(15), partial tetrasomy 15q, or inverted duplication 15 (inv dup 15).
Jacobsen syndrome, which is very rare. It is also called the terminal 11q deletion disorder.[74] Those affected have normal intelligence or mild developmental disability, with poor expressive language skills. Most have a bleeding disorder called Paris-Trousseau syndrome.
Klinefelter syndrome (XXY). Men with Klinefelter syndrome are usually sterile and tend to be taller and have longer arms and legs than their peers. Boys with the syndrome are often shy and quiet and have a higher incidence of speech delay and dyslexia. Without testosterone treatment, some may develop gynecomastia during puberty.
Patau Syndrome, also called D-Syndrome or trisomy-13. Symptoms are somewhat similar to those of trisomy-18, without the characteristic folded hand.
Small supernumerary marker chromosome. This means there is an extra, abnormal chromosome. Features depend on the origin of the extra genetic material. Cat-eye syndrome and isodicentric chromosome 15 syndrome (or Idic15) are both caused by a supernumerary marker chromosome, as is Pallister–Killian syndrome.
Triple-X syndrome (XXX). XXX girls tend to be tall and thin and have a higher incidence of dyslexia.
Turner syndrome (X instead of XX or XY). In Turner syndrome, female sexual characteristics are present but underdeveloped. Females with Turner syndrome often have a short stature, low hairline, abnormal eye features and bone development and a "caved-in" appearance to the chest.
Wolf–Hirschhorn syndrome, which is caused by partial deletion of the short arm of chromosome 4. It is characterized by growth retardation, delayed motor skills development, "Greek Helmet" facial features, and mild to profound mental health problems.
XYY syndrome. XYY boys are usually taller than their siblings. Like XXY boys and XXX girls, they are more likely to have learning difficulties.
Sperm aneuploidy[edit]
Exposure of males to certain lifestyle, environmental and/or occupational hazards may increase the risk of aneuploid spermatozoa.[75] In particular, risk of aneuploidy is increased by tobacco smoking,[76][77] and occupational exposure to benzene,[78] insecticides,[79][80] and perfluorinated compounds.[81] Increased aneuploidy is often associated with increased DNA damage in spermatozoa.
See also[edit]
Aneuploidy
Chromosome segregation
DNA
Genetic deletion
For information about chromosomes in genetic algorithms, see chromosome (genetic algorithm)
Genetic genealogy
Genealogical DNA test
Lampbrush chromosome
List of number of chromosomes of various organisms
Locus (explains gene location nomenclature)
Maternal influence on sex determination
Non-disjunction
Sex-determination system
XY sex-determination system
X-chromosome
X-inactivation
Y-chromosome
Y-chromosomal Aaron
Y-chromosomal Adam
Polytene chromosome
Neochromosome
Parasitic chromosome
Notes and references[edit]
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Jump up ^ Burt DW (2002). "Origin and evolution of avian microchromosomes". Cytogenet. Genome Res. 96 (1–4): 97–112. doi:10.1159/000063018. PMID 12438785.
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Jump up ^ Mendell JE; Clements KD; Choat JH; Angert ER (May 2008). "Extreme polyploidy in a large bacterium". Proc. Natl. Acad. Sci. U.S.A. 105 (18): 6730–4. Bibcode:2008PNAS..105.6730M. doi:10.1073/pnas.0707522105. PMC 2373351  . PMID 18445653.
Jump up ^ White, M. J. D. (1973). The chromosomes (6th ed.). London: Chapman and Hall, distributed by Halsted Press, New York. p. 28. ISBN 0-412-11930-7.
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Jump up ^ Painter, Theophilus S. (April 1923). "Studies in mammalian spermatogenesis. II. The spermatogenesis of man". Journal of Experimental Zoology. 37 (3): 291–336. doi:10.1002/jez.1400370303.
Jump up ^ Tjio JH; Levan A (1956). "The chromosome number of man". Hereditas. 42 (1–2): 1–6. doi:10.1111/j.1601-5223.1956.tb03010.x. hdl:10261/15776.
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Jump up ^ Santaguida, Stefano; Amon, Angelika (2015-08-01). "Short- and long-term effects of chromosome mis-segregation and aneuploidy". Nature Reviews Molecular Cell Biology. 16 (8): 473–485. doi:10.1038/nrm4025. ISSN 1471-0080. PMID 26204159.
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Jump up ^ Xia Y, Bian Q, Xu L, Cheng S, Song L, Liu J, Wu W, Wang S, Wang X (2004). "Genotoxic effects on human spermatozoa among pesticide factory workers exposed to fenvalerate". Toxicology. 203 (1–3): 49–60. doi:10.1016/j.tox.2004.05.018. PMID 15363581.
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Jump up ^ Governini L, Guerranti C, De Leo V, Boschi L, Luddi A, Gori M, Orvieto R, Piomboni P (2015). "Chromosomal aneuploidies and DNA fragmentation of human spermatozoa from patients exposed to perfluorinated compounds". Andrologia. 47 (9): 1012–9. doi:10.1111/and.12371. PMID 25382683.
External links[edit]
Wikimedia Commons has media related to Chromosomes.
An Introduction to DNA and Chromosomes from HOPES: Huntington's Outreach Project for Education at Stanford
Chromosome Abnormalities at AtlasGeneticsOncology
On-line exhibition on chromosomes and genome (SIB)
What Can Our Chromosomes Tell Us?, from the University of Utah's Genetic Science Learning Center
Try making a karyotype yourself, from the University of Utah's Genetic Science Learning Center
Kimballs Chromosome pages
Chromosome News from Genome News Network
Eurochromnet, European network for Rare Chromosome Disorders on the Internet
Ensembl.org, Ensembl project, presenting chromosomes, their genes and syntenic loci graphically via the web
Genographic Project
Home reference on Chromosomes from the U.S. National Library of Medicine
Visualisation of human chromosomes and comparison to other species
Unique - The Rare Chromosome Disorder Support Group Support for people with rare chromosome disorders
[hide]
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Cytogenetics: chromosomes
Basic
concepts
Karyotype
Ploidy
Genetic material/Genome
Chromatin
Euchromatin
Heterochromatin
Chromosome
Chromatid
Nucleosome
Nuclear organization
Types
Autosome/Sex chromosome (or allosome or heterosome)
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Telomere: Telomere-binding protein (TINF2)
Protamine
Histone
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H3
H4
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A
B
C1
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E
F
H
I
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K
M
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T
See also
Extrachromosomal DNA
Plasmid
List of organisms by chromosome count
List of chromosome lengths for various organisms
List of sequenced genomes
International System for Human Cytogenetic Nomenclature
Authority control
GND: 4010162-9
NDL: 00570780
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Bat Masterson (TV series)
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Bat Masterson
Gene Barry as Bat Masterson, 1958
Genre
Western
Starring
Gene Barry
Narrated by
Bill Baldwin, Bob LeMond
Theme music composer
Havens Wray (David D. Rose)
Ending theme
Bill Lee (Singer)
Country of origin
United States
Original language(s)
English
No. of seasons
3
No. of episodes
107
Production
Producer(s)
Frank Pittman
Andy White
Frederick W. Ziv
Running time
30 minutes
Production company(s)
Ziv Television Programs
United Artists Television
Distributor
MGM Television
Peter Rodgers Organization
Release
Original network
NBC
Picture format
Black-and-white
Audio format
Monaural
Original release
October 8, 1958 (1958-10-08) – June 1, 1961 (1961-06-01)
Gene Barry as Bat Masterson (1960)
Gene Barry as Bat Masterson (1958)
Bat Masterson is an American Western television series which showed a fictionalized account of the life of real-life marshal/gambler/dandy Bat Masterson. The title character was played by Gene Barry and the half-hour black-and-white shows ran on NBC from 1958 to 1961. The series was produced by Ziv Television Productions.
Contents
[hide]
1 Overview
2 Cast
2.1 Main cast
2.2 Guest cast
3 Production
3.1 Development
4 Episodes
4.1 Season 1 (37 episodes)
4.2 Season 2 (37 episodes)
4.3 Season 3 (34 episodes)
5 Spin-offs and remakes
5.1 Guns of Paradise (1990)
5.2 The Gambler Returns: The Luck of the Draw (1991)
6 Home Media
7 In other media
7.1 Comics
8 References
9 External links
Overview[edit]
The show took a tongue-in-cheek outlook, with Barry's Masterson often dressed in expensive Eastern clothing and preferring to use his cane rather than a gun to get himself out of trouble, hence the nickname "Bat". Masterson was also portrayed as a ladies' man who traveled the West looking for women and adventure.
The black derby, fancy decorative vest, black jacket, and elegant pearl-tipped cane were his trademarks. Miniaturized versions were marketed to children as tie-in products during the run of the show.[1]
Cast[edit]
Main cast[edit]
Gene Barry as Bat Masterson (108 episodes)
Allison Hayes as Ellie Winters (7 episodes)
Allen Jaffe as Belligerent Drunk (7 episodes)
Ken Drake as Burdette (6 episodes)
Troy Melton as Government Agent (6 episodes)
Guest cast[edit]
Bat Masterson guest stars included the character actor Robert F. Simon, who appeared as Harrison Whitney in the episode "Death by Decree", and Richard Eastham, who appeared in the 1961 episode "A Lesson in Violence". Stephanie Powers appeared in 1961 episode named "Dead Man's Claim" (using the name Taffy Paul) as Ann, the daughter of the boarding house owner. George Macready appeared as Clyde Richards in the 1961 episode "Tempest at Tioga Pass". Quintin Sondergaard appeared in various roles on the series five times between 1958 and 1961. Dan Sheridan was cast as Joe Rankin in the 1959 episode "Election Day".
Yvonne Lime Fedderson was cast as Lola White in the 1960 episode "The Snare". Ron Hayes appeared four times in the role of Wyatt Earp. Gary Vinson appeared as Billy Thompson in the 1958 episode "A Noose Fits Anybody". Tyler McVey appeared from 1958 to 1961 in different roles in three episodes ("Dynamite Blows Two Ways", "Incident at Fort Bowie", and "Dead Man's Claim"). Ron Foster (1960) appeared twice as Toby Dawson in "Six Feet of Gold" and as Sheriff Buck Simpson in "Jeopardy at Jackson Hole" (1961). James Coburn also appeared in "Six Feet of Gold". Tom Greenway guest starred twice, as Charlie in "Buffalo Kill" (1959) and as Ben Pick in "Dagger Dance" (1961). Brett King appeared four times, beginning with the role of Hub Elliott in "License to Cheat" (1959); Douglas Kennedy was cast as Sheriff Jeb Crater in that same episode. William Tannen was cast in four episodes of Bat Masterson.
Diane Brewster, also known as "Beaver" Cleaver's second-grade teacher "Miss Canfield" in Leave It to Beaver and as gambler "Samantha Crawford" in Maverick opposite James Garner and Jack Kelly as well as in a single episode of Cheyenne, played the role of Miss Lynn Harrison in the 2-part episode "The Conspiracy" (1959). Kevin Hagen appeared as Ace Williams in "The Fourth Man" (1961). Lon Chaney, Jr. played the role of Rance Fletcher in "Bat Trap" (1961).
Production[edit]
Development[edit]
The series was loosely based on Richard O'Connor's 1957 biography of Masterson.[2] This was highlighted by the book's front cover being shown at the end of the closing credits with an onscreen notation "based on".
From 1955 to 1959, Mason Alan Dinehart played a 20-something Bat Masterson in thirty-four episodes of the ABC/Desilu western series, The Life and Legend of Wyatt Earp, starring Hugh O'Brian as the frontier peace officer Wyatt Earp. Dinehart left the series, and Barry was hence cast as a 40-something Masterson in a separate series on a different network. In the year 1958-1959 both actors were featured in the role of Masterson.[3]
The theme song was sung by Bill Lee, a member of the Mellomen.
Barry went on to star in Burke's Law, a detective series set in modern-day Los Angeles, as well as The Name of the Game.
Episodes[edit]
The series consisted of three seasons and 107 episodes:
Season 1 (37 episodes)[edit]
With Jean Willes and Adele Mara
Ep #
Title
Airdate
1 "Double Showdown" October 8, 1958
2 "Two Graves for Swan Valley" October 15, 1958
3 "Dynamite Blows Two Ways" October 22, 1958
4 "Stampede at Tent City"
(with James Best and William Conrad) October 29, 1958
5 "The Fighter"
(with Marie Windsor and Robert J. Wilke) November 5, 1958
6 "Bear Bait" November 12, 1958
7 "A Noose Fits Anybody" November 19, 1958
8 "Dude's Folly" November 26, 1958
9 "The Treasure of Worry Hill"
(with Ross Martin) December 3, 1958
10 "Cheyenne Club" December 17, 1958
11 "Sherman's March Through Dodge City" December 24, 1958
12 "Trail Pirate" December 31, 1958
13 "Double Trouble in Trinidad" January 7, 1959
14 "Election Day" January 14, 1959
15 "One Bullet from Broken Bow" January 21, 1959
16 "A Personal Matter" January 8, 1959
17 "License to Cheat" February 4, 1959
18 "Sharpshooter" February 11, 1959
19 "River Boat" February 18, 1959
20 "Battle of the Pass" February 25, 1959
21 "Marked Deck" March 11, 1959
22 "Incident in Leadville" March 18, 1959
23 "The Tumbleweed Wagon" March 25, 1959
24 "Brunette Bombshell" April 1, 1959
25 "Deadline" April 8, 1959
26 "A Matter of Honor"
with John Vivyan April 22, 1959
27 "Man of Action" April 29, 1959
28 "Lottery of Death" May 13, 1959
29 "The Death of Bat Masterson" May 20, 1959
30 "The Secret is Death" May 27, 1959
31 "Promised Land" June 10, 1959
32 "The Conspiracy: Part 1" June 17, 1959
33 "The Conspiracy: Part 2" June 24, 1959
34 "The Black Pearls" July 1, 1959
35 "The Desert Ship" July 15, 1959
36 "The Romany Knives" July 22, 1959
37 "Buffalo Kill" July 29, 1959
Season 2 (37 episodes)[edit]
Ep #
Title
Airdate
1 "To the Manner Born" October 1, 1959
2 "Wanted -- Dead" October 15, 1959
3 "No Funeral for Thorn" October 22, 1959
4 "Shakedown at St. Joe" October 29, 1959
5 "Lady Luck" November 5, 1959
6 "Who'll Bury My Violence?" November 12, 1959
7 "Dead Men Don't Pay Debts" November 19, 1959
8 "Death and Taxes" November 26, 1959
9 "Bat Plays a Dead Man's Hand" December 3, 1959
10 "Garrison Finish" December 10, 1959
11 "The Canvas and the Cane" December 17, 1959
12 "The Inner Circle" December 31, 1959
13 "The Pied Piper of Dodge City" January 7, 1960
14 "A Picture of Death" January 14, 1960
15 "Pigeon and Hawk" January 21, 1960
16 "Flume to the Mother Lode" January 28, 1960
17 "Death by the Half Dozen" February 4, 1960
18 "Deadly Diamonds" February 11, 1960
19 "Mr. Fourpaws" February 18, 1960
20 "Six Feet of Gold" February 25, 1960
21 "Cattle and Cane" March 3, 1960
22 "The Disappearance of Bat Masterson" March 10, 1960
23 "The Snare" March 17, 1960
24 "Three Bullets for Bat" March 24, 1960
25 "The Reluctant Witness" March 31, 1960
26 "Come Out Fighting" April 7, 1960
27 "Stage to Nowhere" April 14, 1960
28 "Incident at Fort Bowie" April 21, 1960
29 "Masterson's Arcadia Club" April 28, 1960
30 "Welcome to Paradise" May 5, 1960
31 "A Grave Situation" May 12, 1960
32 "Gold Is Where You Steal It" May 19, 1960
33 "Wanted -- Alive Please" May 26, 1960
34 "The Elusive Baguette" June 2, 1960
35 "The Big Gamble" June 16, 1960
36 "Blood on the Money" with Walter Coy June 23, 1960
37 "Barbary Castle" July 1, 1960
Season 3 (34 episodes)[edit]
The actual Bat Masterson (1879)
Ep #
Title
Airdate
1 "Debt of Honor" September 29, 1960
2 "Law of the Land" October 6, 1960
3 "Bat Trap" October 13, 1960
4 "The Rage of Princess Ann" October 20, 1960
5 "The Hunter"
with guest star John Vivyan October 27, 1960
6 "Murder Can Be Dangerous" November 3, 1960
7 "High Card Loses" November 10, 1960
8 "Dakota Showdown " November 17, 1960
9 "The Last of the Night Raiders" November 24, 1960
10 "Last Stop to Austin" December 1, 1960
11 "A Time to Die" December 15, 1960
12 "Death by Decree" December 22, 1960
13 "The Lady Plays Her Hand" December 29, 1960
14 "Tempest at Tioga Pass" January 5, 1961
15 "The Court Martial of Major Mars" January 19, 1961
16 "The Price of Paradise" January 19, 1961
17 "End of the Line" January 26, 1961
18 "The Prescott Campaign" February 2, 1961
19 "Bullwhacker's Bounty" February 16, 1961
20 "A Lesson in Violence" February 23, 1961
21 "Run for Your Money" March 2, 1961
22 "Terror on the Trinity" March 9, 1961
23 "Episode in Eden" March 16, 1961
24 "The Good and the Bad" March 23, 1961
25 "No Amnesty for Death"
(with DeForest Kelley) March 30, 1961
26 "Ledger of Guilt" April 6, 1961
27 "Meeting at Mimbres" April 13, 1961
28 "Dagger Dance" April 20, 1961
29 "The Fourth Man" April 27, 1961
30 "Dead Man's Claim" May 4, 1961
31 "The Marble Slab" May 11, 1961
32 "Farmer with a Badge" May 18, 1961
33 "The Fatal Garment" May 25, 1961
34 "Jeopardy at Jackson Hole" June 1, 1961
Spin-offs and remakes[edit]
Guns of Paradise (1990)[edit]
Barry recreated the role of Bat Masterson in an episode of the television series Guns of Paradise (1990), alongside Hugh O'Brian as Wyatt Earp.
The Gambler Returns: The Luck of the Draw (1991)[edit]
In The Gambler Returns: The Luck of the Draw (1991) Barry played Masterson, also with O'Brian as Earp, as well as Jack Kelly as Bart Maverick and Clint Walker as Cheyenne Bodie.[4]
Home Media[edit]
TGG Direct released seasons the first and second seasons on DVD in Region 1 on January 29, 2013.[5][6] The third and final season was released on November 5, 2013.[7] Due to licensing issues, the episode Terror on the Trinity is not included.
In other media[edit]
Comics[edit]
Dell Comics issued nine issues of a quarterly Bat Masterson comic book between Aug./Oct. 1959 and Nov. 1961/Jan. 1962 with the initial issue a Four Color tryout (#1013).[8] Columbia Features syndicated a comic strip from September 7, 1959 to April 1960 written by Ed Herron and drawn by Howard Nostrand (Sept. 1959–Dec. 1959) and Bob Powell (Dec. 1959–April 1960).[9] Notably Nostrand was assisted (on backgrounds) by Neal Adams who had just graduated from the School of Industrial Arts; it was among his first professional art jobs.[10]
References[edit]
Jump up ^ http://sentimental-journeys.com/2010/02/08/bat-masterson-cane--derby-hat--comic-book-ad.aspx
Jump up ^ Robert K. Dearment. Bat Masterson: The Man and the Legend. University of Oklahoma Press , 1989. p.9
Jump up ^ "Full Cast and Crew for The Life and Legend of Wyatt Earp". Internet Movie Data Base. Retrieved July 11, 2013.
Jump up ^ [1]
Jump up ^ Bat Masterson Complete Season One
Jump up ^ Bat Masterson Complete Season Two
Jump up ^ Bat Masterson Season 3
Jump up ^ BAT MASTERSON
Jump up ^ The Badmouths of the West
Jump up ^ Neal Adams: Renaissance Man Part I Archived 2009-11-16 at WebCite
External links[edit]
Wikimedia Commons has media related to Bat Masterson (TV series).
Bat Masterson on IMDb
Bat Masterson at TV.com
Gene Barry Fan Page
Retrieved from "https://en.wikipedia.org/w/index.php?title=Bat_Masterson_(TV_series)&oldid=832193465"
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