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== Callus cells deaths ==
== Callus cells deaths ==
Callus can brown and die during culture, but the causes for callus browning are not well understood. In ''[[Jatropha curcas]]'' callus cells, small organized callus cells became disorganized and varied in size after browning occurred.<ref>{{cite journal|last=He|first=Yang|coauthors=Guo, Xiulian, Lu, Ran, Niu, Bei, Pasapula, Vijaya, Hou, Pei, Cai, Feng, Xu, Ying, Chen, Fang|title=Changes in morphology and biochemical indices in browning callus derived from Jatropha curcas hypocotyls|journal=Plant Cell, Tissue and Organ Culture (PCTOC)|volume=98|issue=1|pages=11–17|doi=10.1007/s11240-009-9533-y}}</ref> [[Browning (chemical process)|Browning]] has also been associated with oxidation and [[Natural phenol|phenolic]] compounds in both explant tissues and explant secretions.<ref>{{cite journal|last=Dan|first=Yinghui|coauthors=Armstrong, Charles L., Dong, Jimmy, Feng, Xiaorong, Fry, Joyce E., Keithly, Greg E., Martinell, Brian J., Roberts, Gail A., Smith, Lori A., Tan, Lalaine J., Duncan, David R.|title=Lipoic acid—an unique plant transformation enhancer|journal=In Vitro Cellular & Developmental Biology - Plant|volume=45|issue=6|pages=630–638|doi=10.1007/s11627-009-9227-5}}</ref>
Callus can brown and die during culture, but the causes for callus browning are not well understood. In ''[[Jatropha curcas]]'' callus cells, small organized callus cells became disorganized and varied in size after browning occurred.<ref>{{cite journal|last=He|first=Yang|coauthors=Guo, Xiulian, Lu, Ran, Niu, Bei, Pasapula, Vijaya, Hou, Pei, Cai, Feng, Xu, Ying, Chen, Fang|title=Changes in morphology and biochemical indices in browning callus derived from Jatropha curcas hypocotyls|journal=Plant Cell, Tissue and Organ Culture (PCTOC)|volume=98|issue=1|pages=11–17|doi=10.1007/s11240-009-9533-y}}</ref> [[Browning (chemical process)|Browning]] has also been associated with oxidation and [[Natural phenol|phenolic]] compounds in both explant tissues and explant secretions.<ref>{{cite journal|last=Dan|first=Yinghui|coauthors=Armstrong, Charles L., Dong, Jimmy, Feng, Xiaorong, Fry, Joyce E., Keithly, Greg E., Martinell, Brian J., Roberts, Gail A., Smith, Lori A., Tan, Lalaine J., Duncan, David R.|title=Lipoic acid—an unique plant transformation enhancer|journal=In Vitro Cellular & Developmental Biology - Plant|volume=45|issue=6|pages=630–638|doi=10.1007/s11627-009-9227-5}}</ref> In rice, presumably, a condition which is favorable for scutellar callus induction induces necrosis too.<ref name="pazuki">{{cite journal |last=Pazuki |first=Arman |last2=Sohani |first2=Mehdi |lastauthoramp=yes |year=2013 |title= Phenotypic evaluation of scutellum-derived calluses in ‘Indica’ rice cultivars |url= http://aas.bf.uni-lj.si/september2013/08Pazuki.pdf |format=PDF |journal= Acta Agriculturae Slovenica |volume=101 |issue=2 |pages=239–247 |doi=10.2478/acas-2013-0020 |accessdate=February 2, 2014}}</ref>


==Uses==
==Uses==

Revision as of 06:55, 3 February 2014

Plant callus (plural calluses or calli) is a mass of unorganized parenchyma cells derived from plant tissue (explants) for use in biological research and biotechnology. In plant biology, callus cells are those cells that cover a plant wound.[1] Callus formation is induced from plant tissues after surface sterilization and plating onto in vitro tissue culture medium. Plant growth regulators, such as auxins, cytokinins, and gibberellins, are supplemented into the medium to initiate callus formation or somatic embryogenesis. Callus initiation has been described for many plant taxonomic divisions:

Callus Nicotiana tabacum

Callus induction and tissue culture

Callus cells forming during a process called "induction" in Pteris vittata

A callus cell culture is usually sustained on gel medium. Callus induction medium consists of agar and a mixture of macronutrients and micronutrients for the given cell type. There are several types of basal salt mixtures used in plant tissue culture, but most notably modified Murashige and Skoog medium,[13] White's medium,[14] and woody plant medium.[15] Vitamins are also provided to enhance growth such as Gamborg B5 vitamins.[16] For plant cells, enrichment with nitrogen, phosphorus, and potassium is especially important.

Callus induced from Pteris vittata gametophytes

Morphology

Plant callus is usually derived from somatic tissues. The tissues used to initiate callus formation depends on plant species and which tissues are available for explant culture. The cells that give rise to callus and somatic embryos usually undergo rapid division or are partially undifferentiated such as meristematic tissue. In alfalfa, Medicago truncatula, however callus and somatic embryos are derived from mesophyll cells that undergo dedifferentiation.[17] Plant hormones are used to initiate callus growth. Specific auxin to cytokinin ratios in plant tissue culture medium give rise to an unorganized growing and dividing mass of callus cells. Callus cultures are often broadly classified as being either compact or friable. Friable calluses fall apart easily, and can be used to generate cell suspension cultures. Callus can directly undergo direct organogenesis and/or embryogenesis where the cells will form an entirely new plant.

Callus cells deaths

Callus can brown and die during culture, but the causes for callus browning are not well understood. In Jatropha curcas callus cells, small organized callus cells became disorganized and varied in size after browning occurred.[18] Browning has also been associated with oxidation and phenolic compounds in both explant tissues and explant secretions.[19] In rice, presumably, a condition which is favorable for scutellar callus induction induces necrosis too.[20]

Uses

Callus cells are not necessarily genetically homogeneous because a callus is often made from structural tissue, not individual cells. Nevertheless, callus cells are often considered similar enough for standard scientific analysis to be performed as if on a single subject. For example, an experiment may have half a callus undergo a treatment as the experimental group, while the other half undergoes a similar but non-active treatment as the control group.

Plant calli can differentiate into a whole plant, a process called regeneration, through addition of plant hormones in culture medium. This ability is known as totipotency. Regeneration of a whole plant from a single cell allows researchers to recover whole plants that have a copy of the transgene in every cell. Regeneration of a whole plant that has some genetically transformed cells and some untransformed cells is called a chimera. In general, chimeras are not useful for genetic research or agricultural applications.

Genes can be inserted into callus cells using biolistic bombardment, also known as a gene gun, or Agrobacterium tumefaciens. Cells that receive the gene of interest can then be recovered into whole plants using a combination of plant hormones. The whole plants that are recovered can be used to experimentally determine gene function(s), or to enhance crop plant traits for modern agriculture.

Callus is of particular use in micropropagation where it can be used to grow genetically identical copies of plants with desirable characteristics.

History

File:Duhameldemonceau.jpg
Engraving of Henri-Louis Duhamel du Monceau by François-Hubert Drouais. He is shown working on his Éléments d’architecture navale, his most famous work. Duhamel du Monceau was the first to describe callus formation that he observed growing over the wound of an elm tree.

Henri-Louis Duhamel du Monceau investigated wound-healing responses in elm trees, and was the first to report formation of callus on live plants.[21]

In 1908, E. F. Simon was able to induce callus from poplar stems that also produced roots and buds.[22] The first reports of callus induction in vitro came from three independent researchers in 1939.[23] P. White induced callus derived from tumor-developing procambial tissues of hybrid Nicotiana glauca that did not require hormone supplementation.[24] Gautheret and Nobecourt were able to maintain callus cultures of carrot using auxin hormone additions.[citation needed]

See also

References

  1. ^ What is Plant Tissue Culture?
  2. ^ Takeda, Reiji. "Growth and sesquiterpenoid production by Calypogeia granulata inoue cells in suspension culture". Planta. 151 (6): 525–530. doi:10.1007/BF00387429. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  3. ^ Peterson, M (2003). "Cinnamic acid 4-hydroxylase from cell cultures of the hornwort Anthoceros agrestis". Planta. 217 (1): 96–101. doi:10.1007/s00425-002-0960-9. PMID 12721853.
  4. ^ Beutelmann, P. (1 January 1977). "Purification and identification of a cytokinin from moss callus cells". Planta. 133 (3): 215–217. doi:10.1007/BF00380679. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  5. ^ Atmane, N. "Histological analysis of indirect somatic embryogenesis in the Marsh clubmoss Lycopodiella inundata (L.) Holub (Pteridophytes)". Plant Science. 156 (2): 159–167. doi:10.1016/S0168-9452(00)00244-2.
  6. ^ Yang, Xuexi. "Hyperaccumulation of arsenic by callus, sporophytes and gametophytes of Pteris vittata cultured in vitro". Plant Cell Reports. 26 (10): 1889–1897. doi:10.1007/s00299-007-0388-6. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  7. ^ Chavez, V. M. "Regeneration of Ceratozamia euryphyllidia (Cycadales, Gymnospermae) plants from embryogenic leaf cultures derived from mature-phase trees". Plant Cell Reports. 17 (8): 612–616. doi:10.1007/s002990050452. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  8. ^ Jeon, MeeHee. "Ginkgolide B production in cultured cells derived from Ginkgo biloba L. leaves". Plant Cell Reports. 14 (8). doi:10.1007/BF00232783. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  9. ^ Finer, John J. (1 January 1989). "Initiation of embryogenic callus and suspension cultures of eastern white pine (Pinus strobus L.)". Plant Cell Reports. 8 (4): 203–206. doi:10.1007/BF00778532. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  10. ^ O'Dowd, Niamh A. "Callus production, suspension culture and in vitro alkaloid yields of Ephedra". Plant Cell, Tissue and Organ Culture. 34 (2): 149–155. doi:10.1007/BF00036095. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  11. ^ Chen, Ying-Chun. "A reliable protocol for plant regeneration from callus culture of Phalaenopsis". In Vitro Cellular & Developmental Biology – Plant. 36 (5): 420–423. doi:10.1007/s11627-000-0076-5. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  12. ^ Burris, Jason N. (10 October 2009). "An Improved Tissue Culture System for Embryogenic Callus Production and Plant Regeneration in Switchgrass (Panicum virgatum L.)". BioEnergy Research. 2 (4): 267–274. doi:10.1007/s12155-009-9048-8. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  13. ^ Murashige, Toshio (July 1962). "A Revised Medium for Rapid Growth and Bio Assays with Tobacco Tissue Cultures". Physiologia Plantarum. 15 (3): 473–497. doi:10.1111/j.1399-3054.1962.tb08052.x. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  14. ^ White, P. R. (Feb 1939). "Potentially unlimited growth of excised plant callus in an artificial nutrient". American Journal of Botany. 26 (2): 59–4. JSTOR 2436709.
  15. ^ Lloyd, G (1981). "Commercially-feasible micropropagation of mountain laurel, Kalmia latifolia, by use of shoot-tip culture". Combined Proceedings, International Plant Propagators' Society. 30: 421–427. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  16. ^ Gamborg, OL (April 1968). "Nutrient requirements of suspension cultures of soybean root cells". Experimental Cell Research. 50 (1): 151–158. doi:10.1016/0014-4827(68)90403-5. PMID 5650857. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  17. ^ Wang, X.-D. (10 January 2011). "Ontogeny of embryogenic callus in Medicago truncatula: the fate of the pluripotent and totipotent stem cells". Annals of Botany. 107 (4): 599–609. doi:10.1093/aob/mcq269. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  18. ^ He, Yang. "Changes in morphology and biochemical indices in browning callus derived from Jatropha curcas hypocotyls". Plant Cell, Tissue and Organ Culture (PCTOC). 98 (1): 11–17. doi:10.1007/s11240-009-9533-y. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  19. ^ Dan, Yinghui. "Lipoic acid—an unique plant transformation enhancer". In Vitro Cellular & Developmental Biology - Plant. 45 (6): 630–638. doi:10.1007/s11627-009-9227-5. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  20. ^ Pazuki, Arman; Sohani, Mehdi (2013). "Phenotypic evaluation of scutellum-derived calluses in 'Indica' rice cultivars" (PDF). Acta Agriculturae Slovenica. 101 (2): 239–247. doi:10.2478/acas-2013-0020. Retrieved February 2, 2014. {{cite journal}}: Unknown parameter |lastauthoramp= ignored (|name-list-style= suggested) (help)
  21. ^ Razdan, M. K. (2003). Introduction to plant tissue culture (2. ed. ed.). Enfield, NH [u.a.]: oxford Publishers. ISBN 1-57808-237-4. {{cite book}}: |edition= has extra text (help)
  22. ^ Gautheret, Roger J. (1 December 1983). "Plant tissue culture: A history". The Botanical Magazine Tokyo. 96 (4): 393–410. doi:10.1007/BF02488184.
  23. ^ Chawla, H.S. (2002). Introduction to plant biotechnology (2nd ed. ed.). Enfield, N.H.: Science Publishers. ISBN 1-57808-228-5. {{cite book}}: |edition= has extra text (help)
  24. ^ White, P. R. (Feb 1939). "Potentially unlimited growth of excised plant callus in an artificial nutrient". American Journal of Botany. 26 (2): 59–4. JSTOR 2436709.