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====Melanoma====
====Melanoma====
Zebrafish expressing mutated forms of either the BRAF or NRAS [[oncogene]]s develop melanoma when placed onto a p53 deficient background. [[Histology|Histologically]], these tumors strongly resemble the human disease, are fully transplantable, and exhibit large-scale genomic alterations. The BRAF melanoma model was utilized as a platform for two screens published in March 2011 in the journal ''Nature''. In one study, by Ceol, Houvras and Zon, the model was used as a tool to understand the functional importance of genes known to be amplified and overexpressed in human melanoma.<ref>{{cite web|url=http://www.ncbi.nlm.nih.gov/pubmed/21430779|title=The histone methyltransferase SETDB1 is recurrently amplified in melanoma and accelerates its onset|publisher=''Nature'' via NCBI|date=March 2011|accessdate=26 October 2012}}</ref> One gene, SETDB1, markedly accelerated tumor formation in the zebrafish system, demonstrating its importance as a new melanoma oncogene. This was particularly significant because SETDB1 is known to be involved in the epigenetic regulation that is increasingly appreciated to be central to tumor cell biology. In another study, by White and Zon, an effort was made to therapeutically target the genetic program present in the tumor's origin [[neural crest]] cell using a chemical screening approach.<ref>{{cite web|url=http://www.ncbi.nlm.nih.gov/pubmed/21430780|title=DHODH modulates transcriptional elongation in the neural crest and melanoma|publisher=''Nature'' via NCBI|date=March 2011|accessdate=26 October 2012}}</ref> This revealed that an inhibition of the DHODH protein (by a small molecule called leflunomide) prevented development of the neural crest stem cells which ultimately give rise to melanoma via interference with the process of transcriptional elongation. Because this approach would aim to target the "identity" of the melanoma cell rather than a single genetic mutation, leflunomide may have utility in treating human melanoma.
Zebrafish expressing mutated forms of either the BRAF or NRAS [[oncogene]]s develop melanoma when placed onto a p53 deficient background. [[Histology|Histologically]], these tumors strongly resemble the human disease, are fully transplantable, and exhibit large-scale genomic alterations. The BRAF melanoma model was utilized as a platform for two screens published in March 2011 in the journal ''Nature''. In one study, by Ceol, Houvras and Zon, the model was used as a tool to understand the functional importance of genes known to be amplified and overexpressed in human melanoma.<ref>{{cite web|url=http://www.ncbi.nlm.nih.gov/pubmed/21430779|title=The histone methyltransferase SETDB1 is recurrently amplified in melanoma and accelerates its onset|publisher=''Nature'' via NCBI|date=March 2011|accessdate=26 October 2012}}</ref> One gene, SETDB1, markedly accelerated tumor formation in the zebrafish system, demonstrating its importance as a new melanoma oncogene. This was particularly significant because SETDB1 is known to be involved in the epigenetic regulation that is increasingly appreciated to be central to tumor cell biology. In another study, by White and Zon, an effort was made to therapeutically target the genetic program present in the tumor's origin [[neural crest]] cell using a chemical screening approach.<ref>{{cite web|url=http://www.ncbi.nlm.nih.gov/pubmed/21430780|title=DHODH modulates transcriptional elongation in the neural crest and melanoma|publisher=''Nature'' via NCBI|date=March 2011|accessdate=26 October 2012}}</ref> This revealed that an inhibition of the DHODH protein (by a small molecule called leflunomide) prevented development of the neural crest stem cells which ultimately give rise to melanoma via interference with the process of transcriptional elongation. Because this approach would aim to target the "identity" of the melanoma cell rather than a single genetic mutation, leflunomide may have utility in my face

===Cardiovascular disease===
===Cardiovascular disease===
In cardiovascular research, the zebrafish has been used to model [[Coagulation|blood clotting]], [[Angiogenesis|blood vessel development]], heart failure, and [[congenital heart defect|congenital heart and kidney disease]].
In cardiovascular research, the zebrafish has been used to model [[Coagulation|blood clotting]], [[Angiogenesis|blood vessel development]], heart failure, and [[congenital heart defect|congenital heart and kidney disease]].

Revision as of 20:28, 6 November 2012

Danio rerio
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
D. rerio
Binomial name
Danio rerio
(F. Hamilton, 1822)
Synonyms
  • Barilius rerio
  • Brachydanio rerio
  • Cyprinus chapalio
  • Cyprinus rerio
  • Danio foankei
  • Danio lineatus
  • Nuria rerio
  • Perilampus striatus

The zebrafish, Danio rerio, is a tropical freshwater fish belonging to the minnow family (Cyprinidae) of order Cypriniformes.[1] It is a popular aquarium fish, frequently sold under the trade name zebra danio, and is an important vertebrate model organism in scientific research. It is particularly notable for its regenerative abilities.[2]

Taxonomy

The zebrafish is a derived member of the genus Danio, of the family Cyprinidae. It has a sister-group relationship with Danio kyathit.[3] Zebrafish are also closely related to the genus Devario, as demonstrated by a phylogenetic tree of close species.[4] The zebrafish was referred to in literature as Brachydanio rerio for many years until its reassignment to the genus Danio.[5]

Distribution

The zebrafish is native to the streams of the southeastern Himalayan region,[3] and is found in parts of India, Pakistan, Bangladesh, Nepal, and Burma.[6] The species arose in the Ganges region in eastern India, and commonly inhabits streams, canals, ditches, ponds, and slow-moving or stagnant water bodies, including rice fields.[7] Zebrafish have been introduced to parts of the United States, presumably by deliberate release or by escape from fish farms.[6]

Description

The zebrafish is named for the five uniform, pigmented, horizontal, blue stripes on the side of the body, which are reminiscent of a zebra's stripes, and which extend to the end of the caudal fin. Its shape is fusiform and laterally compressed, with its mouth directed upwards. The male is torpedo-shaped, with gold stripes between the blue stripes; the female has a larger, whitish belly and silver stripes instead of gold. Adult females will exhibit a small genital papilla in front of the anal fin origin. The zebrafish can grow to 6.4 cm (2.5 in) in length, although it seldom grows larger than 4 cm (1.6 in) in captivity. Its lifespan in captivity is around two to three years, although in ideal conditions, this may be extended to five years.[7]

Reproduction

A juvenile zebrafish.

The approximate generation time for Danio rerio is three to four months. A male must be present for ovulation and spawning to occur. Females are able to spawn at intervals of two to three days, laying hundreds of eggs in each clutch. Upon release, embryonic development begins; absent sperm, growth stops after the first few cell divisions. Fertilized eggs almost immediately become transparent, a characteristic that makes D. rerio a convenient research model species.[7] Development progresses rapidly – precursors to all major organs appear within 36 hours of fertilization, and hatching takes place 12–36 hours later, depending on the embryo's internal conditions and the external temperature, which is ideally 28.5 °C (83.3 °F). Swimming and feeding behavior begin about 36 hours later. The sex of juveniles cannot be distinguished except by dissection, and sex determinants are not clearly understood.

To encourage the fish to spawn, some researchers use a fish tank with a sliding bottom insert, which reduces the depth of the pool. The technique is most effective in the early morning, given that ovulation and spawning are enhanced by light. Researchers have been able to collect 10,000 embryos in 10 minutes using this method.[8] Male zebrafish are furthermore known to respond to more pronounced markings on females, i.e., "good stripes", but in a group, males will mate with whichever females they can find. What attracts females is not currently understood. The presence of plants, even plastic plants, also apparently encourages spawning.[8]

Feeding

The zebrafish is omnivorous. It primarily eats zooplankton, insects, insect larvae, and phytoplankton, although it can eat a variety of other foods, such as worms and small crustaceans, if its preferred sources are not readily available.[7] Most zebrafish accept common food flakes and tubifex worms in the aquarium.

Aquarium care

Zebrafish are hardy fish and considered good for beginner aquarists. Their enduring popularity can be attributed to their playful disposition,[9] as well as their rapid breeding, aesthetics, cheap price, and broad availability. They thrive best in water temperatures of 22–28 °C (72–82 °F), and require an aquarium volume of at least 10 US gallons (38 L). They also do well in schools or shoals of six or more, and interact well with other fish species in the aquarium. However, they are susceptible to Oodinium or velvet disease, microsporidia (Pseudoloma neurophilia), and Mycobacterium species. Given the opportunity, adults eat hatchlings, which may be protected by separating the two groups with a net, breeding box or separate tank.

Strains

In late 2003, transgenic zebrafish that express green, red, and yellow fluorescent proteins became commercially available in the United States. The fluorescent strains are tradenamed GloFish; other cultivated varieties include 'golden', 'sandy', 'longfin' and 'leopard'.

The leopard danio, previously known as Danio frankei, is a spotted colour morph of the zebrafish which arose due to a pigment mutation.[10] Xanthistic forms of both the zebra and leopard pattern, along with long-finned subspecies, have been obtained via selective breeding programs for the aquarium trade.[11]

Wild-type strains

The Zebra Fish Information Network provides up-to-date information about current known wild-type (WT) strains of D. rerio, some of which are listed below.[12]

Hybrids

Hybrids between different Danio species may be fertile: for example, between D. rerio and D. nigrofasciatus.[13][14]

In scientific research

Zebrafish chromatophores, shown here mediating background adaptation, are widely studied by scientists.
A zebrafish pigment mutant (bottom) produced by insertional mutagenesis.[15] A wild-type embryo (top) is shown for comparison. The mutant lacks black pigment in its melanocytes because it is unable to synthesize melanin properly.

D. rerio is a common and useful model organism for studies of vertebrate development and gene function.[3] Its importance has been consolidated by successful large-scale forward genetic screens (commonly referred to as the Tübingen/Boston screens). In 1996, the scholarly journal Development devoted an issue to zebrafish research in celebration of this landmark.[16] The fish has a dedicated online database of genetic, genomic, and developmental information, the Zebrafish Information Network (ZFIN). D. rerio is also one of the few fish species to have been sent into space.

Research with D. rerio has yielded advances in the fields of developmental biology, oncology,[17] toxicology,[18] reproductive studies, teratology, genetics, neurobiology, environmental sciences, stem cell and regenerative medicine,[19] and evolutionary theory.[13]

Model characteristics

As a model biological system, the zebrafish possesses numerous advantages for scientists. Its genome has been fully sequenced, and it has well-understood, easily observable and testable developmental behaviors. Its embryonic development is very rapid, and its embryos are relatively large, robust, and transparent, and able to develop outside their mother.[20] Furthermore, well-characterized mutant strains are readily available.

Other advantages include the species' nearly constant size during early development, which facilitates simple staining techniques, and the fact that its two-celled embryo can be fused into a single cell to create a homozygous embryo. The zebrafish is also demonstrably similar to mammalian models and humans in toxicity testing, and exhibits a diurnal sleep cycle with similarities to mammalian sleep behavior.[21]

Regeneration

Zebrafish have the ability to regenerate their fins, skin, heart and, in larval stages, brain.[22] Zebrafish heart muscle regeneration does not make use of stem cells; instead, mature heart muscle cells regress to a stem cell-like state and redifferentiate.[22] In 2011, the British Heart Foundation ran an advertising campaign publicising their intention to study the applicability of this ability to humans, by "spend[ing] £50 million on a programme of groundbreaking research that could help us begin to repair damaged [human] hearts."[23]

Zebrafish have also been found to regenerate photoreceptor cells and retinal neurons following injury, which has been shown to be mediated by the dedifferentiation and proliferation of Müller glia.[24] Researchers frequently amputate the dorsal and ventral tail fins and analyze their regrowth to test for mutations. It has been found that histone demethylation occurs at the site of the amputation, switching the zebrafish's cells to an "active", regenerative, stem cell-like state.[25] In 2012, Australian scientists published a study revealing that zebrafish use a specialised protein, known as fibroblast growth factor, to ensure their spinal cords heal without glial scarring after injury.[2]

Another focus of zebrafish research is to understand how a gene called Hedgehog, a biological signal that underlies a number of human cancers, controls cell growth. In probing disorders of the nervous system, including neurodegenerative diseases, movement disorders, psychiatric disorders and deafness, researchers are using the zebrafish to understand how the genetic defects underlying these conditions cause functional abnormalities in the human brain, spinal cord and sensory organs. Researchers are also delving into the complexities of muscle degeneration in genetic models of human musculoskeletal diseases, such as muscular dystrophy.

Genetics

Gene expression

Due to their short lifecycles and relatively large clutch sizes, zebrafish are a useful model for genetic studies. A common reverse genetics technique is to reduce gene expression or modify splicing using Morpholino antisense technology. Morpholino oligonucleotides (MO) are stable, synthetic macromolecules that contain the same bases as DNA or RNA; by binding to complementary RNA sequences, they reduce the expression of specific genes. The journal Genesis[26] devoted an issue[27] to research using MO, mostly in D. rerio. MO can be injected into one cell of an embryo after the 32-cell stage, reducing gene expression in only cells descended from that cell. However, cells in the early embryo (less than 32 cells) are interpermeable to large molecules,[28][29] allowing diffusion between cells. A known problem with gene knockdowns is that, because the genome underwent a duplication after the divergence of ray-finned fishes and lobe-finned fishes, it is not always easy to silence the activity one of the two gene paralogs reliably due to complementation by the other paralog.

Despite the complications of the zebrafish genome, a number of commercially available global platforms exist for analysis of both gene expression by microarrays and promoter regulation using ChIP-on-chip.

Genome sequencing

The Wellcome Trust Sanger Institute started the zebrafish genome sequencing project in 2001, and the full genome sequence of the Tuebingen reference strain is publicly available at the National Center for Biotechnology Information (NCBI)'s Zebrafish Genome Page. The zebrafish reference genome sequence is annotated as part of the Ensembl project, and is maintained by the Genome Reference Consortium.[30]

In 2009, researchers at the Institute of Genomics and Integrative Biology in Delhi announced the sequencing of the genome of a wild zebrafish strain, containing 1.7 billion genetic letters.[31][32]

Mitochondrial DNA

In October 2001, researchers from the University of Oklahoma published D. rerio's complete mitochondrial DNA sequence.[33] Its length is 16,596 base pairs. This is within 100 base pairs of other related species of fish, and it is notably only 18 pairs longer than the goldfish (Carassius auratus) and 21 longer than the carp (Cyprinus carpio). Its gene order and content are identical to the common vertebrate form of mitochondrial DNA. It contains 13 protein-coding genes and a noncoding control region containing the origin of replication for the heavy strand. In between a grouping of five tRNA genes, a sequence resembling vertebrate origin of light strand replication is found. It is difficult to draw evolutionary conclusions because it is difficult to determine whether base pair changes have adaptive significance via comparisons with other vertebrates nucleotide sequences.

Pigmentation genes

In December 2005, a study of the golden strain identified the gene responsible for its unusual pigmentation as SLC24A5, a solute carrier that appeared to be required for melanin production, and confirmed its function with a Morpholino knockdown. The orthologous gene was then characterized in humans and a one base pair difference was found to strongly segregate fair-skinned Europeans and dark-skinned Africans.[34] In developing embryos, melanin production begins at approximately 24 hours post fertilization (hpf), and can be inhibited by treatment with propylthiouracil (PTU).

Transgenesis

Transgenesis is a popular approach to study the function of genes in zebrafish. Construction of transgenic zebrafish is rather easy by a method using the Tol2 transposon system.[35]

Transparent adult bodies

In 2008, researchers at Boston Children's Hospital developed a new strain of zebrafish, named Casper, whose adult bodies were transparent.[36] This allows for detailed visualization of cellular activity, circulation, metastasis and many other phenomena. Because many gene functions are shared between fish and humans, Casper is expected to yield insight into human diseases such as leukemia and other cancers.[36][37]

Use in environmental monitoring

In January 2007, Chinese researchers at Fudan University genetically modified zebrafish to detect oestrogen pollution in lakes and rivers, which is linked to male infertility.[38]

In medical research

Cancer

Zebrafish have been used to make several transgenic models of cancer, including melanoma, leukemia, pancreatic cancer and hepatocellular carcinoma. The flexibility of the fish in transgenesis and for studying genetic interactions, added to its optical properties, make it a unique model for the study of in vivo cancer biology.

Melanoma

Zebrafish expressing mutated forms of either the BRAF or NRAS oncogenes develop melanoma when placed onto a p53 deficient background. Histologically, these tumors strongly resemble the human disease, are fully transplantable, and exhibit large-scale genomic alterations. The BRAF melanoma model was utilized as a platform for two screens published in March 2011 in the journal Nature. In one study, by Ceol, Houvras and Zon, the model was used as a tool to understand the functional importance of genes known to be amplified and overexpressed in human melanoma.[39] One gene, SETDB1, markedly accelerated tumor formation in the zebrafish system, demonstrating its importance as a new melanoma oncogene. This was particularly significant because SETDB1 is known to be involved in the epigenetic regulation that is increasingly appreciated to be central to tumor cell biology. In another study, by White and Zon, an effort was made to therapeutically target the genetic program present in the tumor's origin neural crest cell using a chemical screening approach.[40] This revealed that an inhibition of the DHODH protein (by a small molecule called leflunomide) prevented development of the neural crest stem cells which ultimately give rise to melanoma via interference with the process of transcriptional elongation. Because this approach would aim to target the "identity" of the melanoma cell rather than a single genetic mutation, leflunomide may have utility in my face

Cardiovascular disease

In cardiovascular research, the zebrafish has been used to model blood clotting, blood vessel development, heart failure, and congenital heart and kidney disease.

Immune system

In programmes of research into acute inflammation, a major underpinning process in many diseases, researchers have established a zebrafish model of inflammation, and its resolution. This approach allows detailed study of the genetic controls of inflammation and the possibility of identifying potential new drugs.

Repairing retinal damage

Another notable characteristic of the zebrafish is that it possesses four types of cone cell, with ultraviolet supplementing the red, green and blue cone cell subtypes found in humans. Zebrafish can thus observe a vast spectrum of colours. The species is also studied to better understand the development of the retina; in particular, how the cone cells of the retina become arranged into the so-called 'cone mosaic'. Zebrafish, in addition to certain other teleost fish, are particularly noted for having extreme precision of cone cell arrangement.[41]

This study of the zebrafish's retinal characteristics has also extrapolated into medical enquiry. In 2007, researchers at University College London grew a type of zebrafish adult stem cell found in the eyes of fish and mammals that develops into neurons in the retina. These s could be injected into the eye to treat diseases that damage retinal neurons—nearly every disease of the eye, including macular degeneration, glaucoma, and diabetes-related blindness. The researchers studied Müller glial cells in the eyes of humans aged from 18 months to 91 years, and were able to develop them into all types of retinal neurons. They were also able to grow them easily in the lab. The stem cells successfully migrated into diseased rats' retinas, and took on the characteristics of the surrounding neurons. The team is working to develop the same approach in humans.[42]

Drug discovery

As demonstrated through ongoing research programmes, the zebrafish model affords an ideal opportunity, not only to identify novel candidates for genes underlying human disease, but also to develop novel therapeutic agents in drug discovery programmes.[43] Zebrafish embryos have proven to be a rapid, cost-efficient, and reliable teratology assay model.[44]

See also

References

  1. ^ Froese, Rainer; Pauly, Daniel (eds.) (2007). "Danio rerio" in FishBase. March 2007 version.
  2. ^ a b "Study Reveals Secret of Zebrafish". Sci-News.com. June 1, 2012. Retrieved June 2, 2012.
  3. ^ a b c Mayden, Richard L. (2007). "Phylogenetic relationships of Danio within the order Cypriniformes: a framework for comparative and evolutionary studies of a model species". J. Exp. Zool. (Mol. Dev. Evol.). 308B (5): 642–654. doi:10.1002/jez.b.21175. PMID 17554749. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  4. ^ Heredity. "Figure 2. Phylogenetic relationships among ''danios''". Nature.com. Retrieved July 22, 2012.
  5. ^ "The zebrafish book". ZFIN. Retrieved July 15, 2012.
  6. ^ a b "USGS NAS – Nonindigenous Aquatic Species". Nas.er.usgs.gov. July 22, 2003. Retrieved July 22, 2012.
  7. ^ a b c d Spence R, Gerlach G, Lawrence C, Smith C (2008). "The behaviour and ecology of the zebrafish, Danio rerio". Biological Reviews. 83 (1): 13–34. doi:10.1111/j.1469-185X.2007.00030.x. PMID 18093234. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  8. ^ a b Dockser, Amy (January 13, 2012). "Birds Do It, Bees Do It, Even Zebrafish Do It—Just Too Little". The Wall Street Journal. Retrieved February 11, 2012.
  9. ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1046/j.1474-9728.2002.00012.x, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1046/j.1474-9728.2002.00012.x instead.
  10. ^ Watanabe M; Iwashita M; Ishii M; et al. (2006). "Spot pattern of leopard Danio is caused by mutation in the zebrafish connexin41.8 gene". EMBO Rep. 7 (9): 893–7. doi:10.1038/sj.embor.7400757. PMC 1559663. PMID 16845369. {{cite journal}}: Unknown parameter |author-separator= ignored (help); Unknown parameter |month= ignored (help)
  11. ^ Mills, Dick (1993). Eyewitness Hnbk Aquarium Fish. Harper Collins. ISBN 0-7322-5012-9.
  12. ^ "ZFIN". ZFIN. Retrieved July 22, 2012.
  13. ^ a b Parichy, DM (2006). "Evolution of danio pigment pattern development". Heredity. 97 (3): 200–210. doi:10.1038/sj.hdy.6800867. PMID 16835593. {{cite journal}}: Unknown parameter |month= ignored (help)
  14. ^ Heredity. "Figure 1. Danio pigment pattern diversity and phenotypes of D. rerio hybrids with other danios. Photos". Nature.com. Retrieved July 22, 2012.
  15. ^ See link for pigmentation mutants of D rerio: Figure 5. Pigment pattern mutants within D. rerio. Mutant names are shown along with gene identities in parentheses when known. For example, the picasso phenotype results from mutations in errb3
  16. ^ <Please add first missing authors to populate metadata.> (1996). "Zebrafish issue". Development. 123 (1). {{cite journal}}: Unknown parameter |month= ignored (help)
  17. ^ Xiang J; et al. (2009). Isalan, Mark (ed.). "Identifying Tumor Cell Growth Inhibitors by Combinatorial Chemistry and Zebrafish Assays". PLoS ONE. 4 (2): e4361. doi:10.1371/journal.pone.0004361. PMC 2633036. PMID 19194508. {{cite journal}}: Unknown parameter |author-separator= ignored (help); Unknown parameter |month= ignored (help)CS1 maint: unflagged free DOI (link)
  18. ^ Hill AJ, Teraoka H, Heideman W & Peterson RE (2005). "Zebrafish as a Model Vertebrate for Investigating Chemical Toxicity". Toxicological Sciences. 86 (1): 6–19. doi:10.1093/toxsci/kfi110. PMID 15703261. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  19. ^ Major RJ, Poss KD (2007). "Zebrafish Heart Regeneration as a Model for Cardiac Tissue Repair". Drug Discov Today Dis Models. 4 (4): 219–225. doi:10.1016/j.ddmod.2007.09.002. PMC 2597874. PMID 19081827.
  20. ^ Dahm, Ralf (2006). "The Zebrafish Exposed". American Scientist. 94 (5): 446–453.
  21. ^ Jones, Rachel (October 16, 2007). "Let Sleeping Zebrafish Lie: A New Model for Sleep Studies". PLoS Biology. Vol. 5, no. 10. Public Library of Science. p. e281. doi:10.1371/journal.pbio.0050281. PMC 2020498. PMID 20076649. Retrieved March 27, 2011.{{cite news}}: CS1 maint: unflagged free DOI (link)
  22. ^ a b Wade, Nicholas (March 24, 2010). "Research Offers Clue Into How Hearts Can Regenerate in Some Species". New York Times.
  23. ^ "British Heart Foundation – The science behind the appeal". Bhf.org.uk. February 16, 2007. Retrieved February 11, 2012.
  24. ^ Template:Cite PMID
  25. ^ "Organ Regeneration In Zebrafish: Unraveling The Mechanisms". Science Daily. November 2, 2009. Retrieved June 2, 2012.
  26. ^ "genesis – The Journal of Genetics and Development". .interscience.wiley.com. Retrieved July 22, 2012.
  27. ^ <Please add first missing authors to populate metadata.> (2001). "Zebrafish issue". Genesis. 30 (3). {{cite journal}}: Unknown parameter |month= ignored (help)
  28. ^ Kimmel CB, Law RD (1985). "Cell lineage of zebrafish blastomeres. I. Cleavage pattern and cytoplasmic bridges between cells". Dev Biol. 108 (1): 78–85. doi:10.1016/0012-1606(85)90010-7. PMID 3972182. {{cite journal}}: Unknown parameter |month= ignored (help)
  29. ^ Kimmel CB, Law RD (1985). "Cell lineage of zebrafish blastomeres. III. Clonal analyses of the blastula and gastrula stages". Dev Biol. 108 (1): 94–101. doi:10.1016/0012-1606(85)90012-0. PMID 3972184. {{cite journal}}: Unknown parameter |month= ignored (help)
  30. ^ GRC. "Genome Reference Consortium". GRC. Retrieved October 23, 2012.
  31. ^ Decoding the Genome Mystery. Indian Express, July 5, 2009.
  32. ^ FishMap Zv8. Institute of Genomics and Integrative Biology (IGIB). Retrieved June 7, 2012.
  33. ^ Broughton RE, Milam JE, Roe BA (2001). "The Complete Sequence of the Zebrafish (Danio rerio) Mitochondrial Genome and Evolutionary Patterns in Vertebrate Mitochondrial DNA". Genome Res. 11 (11): 1958–67. doi:10.1101/gr.156801. PMC 311132. PMID 11691861. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  34. ^ Lamason RL; Mohideen MA; Mest JR; et al. (2005). "SLC24A5, a putative cation exchanger, affects pigmentation in zebrafish and humans". Science. 310 (5755): 1782–6. doi:10.1126/science.1116238. PMID 16357253. {{cite journal}}: Unknown parameter |author-separator= ignored (help); Unknown parameter |month= ignored (help)
  35. ^ Kawakami, K; et al. (2004). "A transposon-mediated gene trap approach identifies developmentally regulated genes in zebrafish". Developmental Cell. 7 (1): 133–144. doi:10.1016/j.devcel.2004.06.005. PMID 15239961.
  36. ^ a b White RM; Sessa A; Burke C; et al. (2008). "Transparent adult zebrafish as a tool for in vivo transplantation analysis". Cell Stem Cell. 2 (2): 183–9. doi:10.1016/j.stem.2007.11.002. PMC 2292119. PMID 18371439. {{cite journal}}: Unknown parameter |author-separator= ignored (help); Unknown parameter |month= ignored (help)
  37. ^ "Scientists Create See-Through Fish, Watch Cancer Grow". LiveScience. February 6, 2008. Retrieved July 22, 2012.
  38. ^ Song Houyan and Zhong Tao, professors at Fudan's molecular medicine lab, spent three years cloning estrogen-sensitive genes and injecting them into the fertile eggs of zebrafish. The modified fish turn green if they are placed into water that is polluted by estrogen. Fudan scientists turn fish into estrogen alerts
  39. ^ "The histone methyltransferase SETDB1 is recurrently amplified in melanoma and accelerates its onset". Nature via NCBI. March 2011. Retrieved October 26, 2012. {{cite web}}: Italic or bold markup not allowed in: |publisher= (help)
  40. ^ "DHODH modulates transcriptional elongation in the neural crest and melanoma". Nature via NCBI. March 2011. Retrieved October 26, 2012. {{cite web}}: Italic or bold markup not allowed in: |publisher= (help)
  41. ^ "Ontogeny of cone photoreceptor mosaics in zebrafish - Allison - 2010 - The Journal of Comparative Neurology". Wiley Online Library. June 17, 2010. doi:10.1002/cne.22447. Retrieved July 22, 2012. {{cite web}}: Check |doi= value (help)
  42. ^ Zebra fish may point way to cure for blindness The China Post Friday, August 3, 2007.
  43. ^ "Fish for Science". University of Sheffield. 2011. Retrieved March 19, 2011.
  44. ^ Brannen, Kimberly C. (February 1, 2010). "Development of a zebrafish embryo teratogenicity assay and quantitative prediction model". Birth Defects Research Part B: Developmental and Reproductive Toxicology. 89 (1): 66–77. doi:10.1002/bdrb.20223. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

Further reading