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Polytene chromosomes in a Chironomus salivary gland cell

Polytene chromosome

Polytene chromosomes are oversized chromosomes which have developed from standard chromosomes and are commonly found in the salivary glands of Drosophila melanogaster. Specialized cells undergo repeated rounds of DNA replication without cell division (endomitosis), to increase cell volume, forming a giant polytene chromosome. Polytene chromosomes form when multiple rounds of replication produce many sister chromatids that remain fused together.

Function

In addition to increasing the volume of the cells' nuclei and causing cell expansion, polytene cells may also have a metabolic advantage as multiple copies of genes permits a high level of gene expression. In Drosophila melanogaster, for example, the chromosomes of the larval salivary glands undergo many rounds of endoreduplication, to produce large amounts of glue before pupation. Another example within the organism itself is the tandem duplication of various polytene bands located near the centromere of the X chromosome which results in the Bar phenotype of kidney-shaped eyes.^[1]

Structure

Polytene chromosomes were discovered by Balbiani in 1881. They are commonly found in the salivary glands of insects. Hence it is also called salivary gland chromosome. It is larger in size. Hence it is a giant chromosome. The larger size of the chromosome is due to the presence of many longitudinal strands called chromonemata. Hence named as polytene chromosomes (many stranded). The many strands of the giant chromosome is due to repeated division of the chromosome without the cytoplasmic division. This type of division is called endomitosis. The polytene chromosome contains two types of bands namely dark bands and inters bands. The dark bands are darkly stained and the inter bands are lightly stained with nuclear stains. The dark bands contain more DNA and less RNA. The inter bands contain more RNA and less DNA. The bands of polytene chromosomes become enlarged at certain times to form swellings called puffs. The formation of puffs is called puffing. In the regions of puffs, the chromonemata uncoil and open out to form many loops. The puffing is caused by the uncoiling of individual chromomeres in a band. The puffs indicate the site of active genes when mRNA synthesis takes place. The chromonemata of puff give out many series of loops laterally. As these loops appear as rings, they are called Balbiani rings after the name of the researcher who discovered them. They are formed of DNA, RNA and a few proteins. Lamp brush Chromosome was discovered by Ruckert in 1892. It is a large chromosome. It contains lateral loops and appears like a brush. Hence the name lamp brush chromosome. It is found in the oocytes of Sagitta, Sepia, Echinaster (Echinoderm), insects, sharks, amphibians, reptiles and birds. It is 5900 µm in size. Hence, it is called a giant chromosome. Each lamp brush chromosome consists of main axis and many lateral loops. The main axis of each chromosome is formed of 4 chromatids. The main axis contains a series of thickenings called chromomeres. From each chromomere a pair of lateral loops one on each side arises. Each loop has an axial fibre. The axial fibre is the continuation of the chromonema of the main axis. It is rich in DNA. The axial fibre of the loop is surrounded by a matrix. The matrix is formed of RNA and proteins. The matrix gives a fuzzy appearance. The synthesis of protein and yolk takes place in the lateral loops. These proteins are being used in the early development of embryos. <Arumugam, N.2011. Cell & Molecular Biology, Saras Publication 268-270 >

History

Polytene chromosomes were originally observed in the larval salivary glands of Chironomus midges by Balbiani in 1881,^[2] but the hereditary nature of these structures was not confirmed until they were studied in Drosophila melanogaster in the early 1930s by Emil Heitz and Hans Bauer. They are known to occur in secretory tissues of other dipteran insects such as the Malpighian tubules of Sciara and also in protists, plants, mammals, or in cells from other insects. Some of the largest polytene chromosomes described thus far (see scale bar in figure below) occur in larval salivary gland cells of the Chironomid genus Axarus.

Polytene chromosomes are also used to identify the species of Chironomid larvae that are notoriously difficult to identify. Each morphologically distinct group of larvae consists of a number of morphologically identical (sibling) species that can only be identified by rearing adult males or by cytogenetic analysis of the polytene chromosomes of the larvae. Karyotypes are used to confirm the presence of specific species and to study genetic diversity in species with a wide range.^[3]^[4]

References

^ Hartwell, Leland; Leroy Hood; Michael L. Goldberg; Ann E. Reynolds; Lee M. Silver (2011). Genetics:From Genes to Genomes; Fourth Edition. New York, NY: McGraw-Hill. ISBN 978-0-07-352526-6.
^ Balbiani EG (1881). "Sur la structure du noyau des cellules salivaires chez les larves de Chironomus". Zool. Anz. 4: 637–641.
^ Int Panis L, Kiknadze I, Bervoets L, Aimanova A (1994). "Karyological identification of some species of the genus Chironomus Meigen, 1803 from Belgium". Bull. Annls Soc. R. Belge Ent. 130: 135–142.
^ Кикнадзе ИИ; Михайлова П; Истомина АГ; Голыгина ВВ; Инт Панис Л; Крастанов Б (2006). "Хромосомный полиморфизм и дивергенция популяций у Chironomus nuditarsis Keyl (Diptera, Chironomidae)". Tsitologia. 48: 595–609.

Other references

Baudisch W (1977). "Balbiani ring pattern and biochemical activities in the salivary gland of Acricotopus lucidus (Chironomidae)". Results Probl Cell Differ. 8: 197–212. PMID 335467.
Bridges CB (1935). "Salivary chromosome maps with a key to the banding of the chromosomes of Drosophila melanogaster". J Heredity. 26: 60–64.
Daneholt B (1992). "The transcribed template and the transcription loop in Balbiani rings". Cell Biol Int Rep. 16 (8): 709–715. PMID 1446347. doi:10.1016/S0309-1651(05)80015-3.
Werle SF, E Klekowski & DG Smith (2004). "Inversion polymorphism in a Connecticut River Axarus species (Diptera: Chironomidae): biometric effects of a triple inversion heterozygote". Can. J. Zool. 82: 118–129. doi:10.1139/z03-227.
Pavan, C. and Breuer, M. E. (1952). "Polytene chromosomes in different tissues of Rhynchosciara". Journal of Heredity. 63: 151–157.
Pavan, C. (1967). "Chromosomal changes induced by infective agents Triangle". Sandoz J. Med. Sci. 8 (2): 42–48. PMID 5602878.
Pavan, C., Biesele, J., Riess, R. W. and Wertz, A. V. (1971). "XIII. Changes in the ultrastructure of Rhynchosciara cells infected by Microsporidia". Studies in Genetics. VI: 7103.
Pavan, C., Da Cunha, A. B. and Morsoletto, C. (1971). "Virus-chromosome relationships in cells of Rhynchosciara (Diptera, Sciaridae)". Caryologia. 24 (3): 371–389. doi:10.1080/00087114.1971.10796445.

External links

[1] (87A&C Heat-shock puffs)
[2] (High resolution spreads)
Phaseolus Polytene chromosomes (Plants)

English Journal

Treasure Your Exceptions: An Interview with 2017 George Beadle Award Recipient Susan A. Gerbi.

Gerbi SA1.
Genetics.Genetics.2017 Dec;207(4):1215-1217. doi: 10.1534/genetics.117.300398.
PMID 29203696

Morphology and Polytene Chromosomes of a New Species of Simulium (Trichodagmia) (Diptera: Simuliidae) from the Espinhaço Mountains of Brazil.

do Nascimento JMC1, Hamada N1, Adler PH2.
Journal of medical entomology.J Med Entomol.2017 Nov 29. doi: 10.1093/jme/tjx195. [Epub ahead of print]
PMID 29194522

Polytene Chromosome Structure and Somatic Genome Instability.

Spradling AC1.
Cold Spring Harbor symposia on quantitative biology.Cold Spring Harb Symp Quant Biol.2017 Nov 22. pii: 033670. doi: 10.1101/sqb.2017.82.033670. [Epub ahead of print]
PMID 29167281

Japanese Journal

The structural and molecular analysis of endoreduplicated nuclei in tomato (Solanum lycopersicum) fruit provides evidence for a ploidy-dependent increase in transcriptional activity

Pirrello Julien,Bourdon Matthieu,Cheniclet Catherine [他],Coriton Olivier,Bourge Mickaël,Brown Spencer,Renaudin Jean-Pierre,Frangne Nathalie,Chevalier Christian
Plant Biotechnology 30(3), 301-307, 2013
… The use of the DNA FISH method demonstrated that endopolyploidy leads to the formation of polytene chromosomes, whereas the use of a RNA FISH method demonstrated that the rDNA transcription was increased during polyploidization. …
NAID 130003362435