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Polyploid cells and organisms are those containing more than two paired (homologous) sets of chromosomes. Most species whose cells have nuclei (Eukaryotes) are diploid, meaning they have two sets of chromosomes—one set inherited from each parent. However, polyploidy is found in some organisms and is especially common in plants. In addition, polyploidy also occurs in some tissues of animals that are otherwise diploid, such as human muscle tissues.[1] This is known as endopolyploidy. Species whose cells do not have nuclei (Prokaryotes) are haploid organisms, with only a single chromosome in each cell. Most eukaryotes have diploid cells, but produce haploid gametes (eggs and sperm) by meiosis. In plants and multicellular algae the gametophyte generation is haploid, and produces gametes by mitosis. A monoploid has only one set of chromosomes, and the term is usually only applied to cells or organisms that are normally diploid. Male bees and other Hymenoptera, for example, are monoploid.
Polyploidy refers to a numerical change in a whole set of chromosomes. Organisms in which a particular chromosome, or chromosome segment, is under- or overrepresented are said to be aneuploid (from the Greek words meaning "not", "good", and "fold"). Therefore the distinction between aneuploidy and polyploidy is that aneuploidy refers to a numerical change in part of the chromosome set, whereas polyploidy refers to a numerical change in the whole set of chromosomes.[2]
Polyploidy may occur due to abnormal cell division, either during mitosis, or commonly during metaphase I in meiosis.
Polyploidy occurs in some animals, such as goldfish,[3] salmon, and salamanders, but is especially common among ferns and flowering plants (see Hibiscus rosa-sinensis), including both wild and cultivated species. Wheat, for example, after millennia of hybridization and modification by humans, has strains that are diploid (two sets of chromosomes), tetraploid (four sets of chromosomes) with the common name of durum or macaroni wheat, and hexaploid (six sets of chromosomes) with the common name of bread wheat. Many agriculturally important plants of the genus Brassica are also tetraploids. Polyploidization is a mechanism of sympatric speciation because polyploids are usually unable to interbreed with their diploid ancestors.
Polyploidy can be induced in plants and cell cultures by some chemicals: the best known is colchicine, which can result in chromosome doubling, though its use may have other less obvious consequences as well. Oryzalin will also double the existing chromosome content.
Polyploid types are labeled according to the number of chromosome sets in the nucleus:
- triploid (three sets; 3x), for example seedless watermelons, common in the phylum Tardigrada[4]
- tetraploid (four sets; 4x), for example Salmonidae fish,[5] the cotton Gossypium hirsutum [6]
- pentaploid (five sets; 5x), for example Kenai Birch (Betula papyrifera var. kenaica)
- hexaploid (six sets; 6x), for example wheat, kiwifruit[7]
- heptaploid or septaploid (seven sets; 7x)
- octaploid or octoploid, (eight sets; 8x), for example Acipenser (genus of sturgeon fish), dahlias
- decaploid (ten sets; 10x), for example certain strawberries
- dodecaploid (twelve sets; 12x), for example the plant Celosia argentea or the invasive one Spartina anglica[8] or the amphibian Xenopus ruwenzoriensis.
Examples in animals are more common in non-vertebrates[9] such as flatworms, leeches, and brine shrimp. Within vertebrates, examples of stable polyploidy include the salmonids and many cyprinids (i.e. carp).[10] Some fish have as many as 400 chromosomes.[10] Polyploidy also occurs commonly in amphibians; for example the biomedically-important Xenopus genus contains many different species with as many as 12 sets of chromosomes (dodecaploid).[11] Polyploid lizards are also quite common, but are sterile and must reproduce by parthenogenesis.[citation needed] Polyploid mole salamanders (mostly triploids) are all female and reproduce by kleptogenesis,[12] "stealing" spermatophores from diploid males of related species to trigger egg development but not incorporating the males' DNA into the offspring. While mammalian liver cells are polyploid, rare instances of polyploid mammals are known, but most often result in prenatal death.
An octodontid rodent of Argentina's harsh desert regions, known as the Plains Viscacha-Rat (Tympanoctomys barrerae) has been reported as an exception to this 'rule'.[13] However, careful analysis using chromosome paints shows that there are only two copies of each chromosome in T. barrerae, not the four expected if it were truly a tetraploid.[14] The rodent is not a rat, but kin to guinea pigs and chinchillas. Its "new" diploid [2n] number is 102 and so its cells are roughly twice normal size. Its closest living relation is Octomys mimax, the Andean Viscacha-Rat of the same family, whose 2n = 56. It was therefore surmised that an Octomys-like ancestor produced tetraploid (i.e., 2n = 4x = 112) offspring that were, by virtue of their doubled chromosomes, reproductively isolated from their parents.
Polyploidy was induced in fish by Har Swarup (1956) using a cold-shock treatment of the eggs close to the time of fertilization, which produced triploid embryos that successfully matured.[15][16] Cold or heat shock has also been shown to result in unreduced amphibian gametes, though this occurs more commonly in eggs than in sperm.[17] John Gurdon (1958) transplanted intact nuclei from somatic cells to produce diploid eggs in the frog, Xenopus (an extension of the work of Briggs and King in 1952) that were able to develop to the tadpole stage.[18] The British Scientist, J. B. S. Haldane hailed the work for its potential medical applications and, in describing the results, became one of the first to use the word "clone" in reference to animals. Later work by Shinya Yamanaka showed how mature cells can be reprogrammed to become pluripotent, extending the possibilities to non-stem cells. Gurdon and Yamanaka were jointly awarded the Nobel Prize in 2012 for this work.[18]
True polyploidy rarely occurs in humans, although polyploid cells occur in highly differentiated tissue, such as liver parenchyma and heart muscle, and in bone marrow.[19] Aneuploidy is more common.
Polyploidy occurs in humans in the form of triploidy, with 69 chromosomes (sometimes called 69,XXX), and tetraploidy with 92 chromosomes (sometimes called 92,XXXX). Triploidy, usually due to polyspermy, occurs in about 2–3% of all human pregnancies and ~15% of miscarriages.[citation needed] The vast majority of triploid conceptions end as a miscarriage; those that do survive to term typically die shortly after birth. In some cases, survival past birth may extend longer if there is mixoploidy with both a diploid and a triploid cell population present.
Triploidy may be the result of either digyny (the extra haploid set is from the mother) or diandry (the extra haploid set is from the father). Diandry is mostly caused by reduplication of the paternal haploid set from a single sperm, but may also be the consequence of dispermic (two sperm) fertilization of the egg.[20] Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid oocyte or failure to extrude one polar body from the oocyte. Diandry appears to predominate among early miscarriages, while digyny predominates among triploidy that survives into the fetal period.[citation needed] However, among early miscarriages, digyny is also more common in those cases <8.5 weeks gestational age or those in which an embryo is present. There are also two distinct phenotypes in triploid placentas and fetuses that are dependent on the origin of the extra haploid set. In digyny, there is typically an asymmetric poorly grown fetus, with marked adrenal hypoplasia and a very small placenta.[citation needed] In diandry, a partial hydatidiform mole develops.[20] These parent-of-origin effects reflect the effects of genomic imprinting.[citation needed]
Complete tetraploidy is more rarely diagnosed than triploidy, but is observed in 1–2% of early miscarriages. However, some tetraploid cells are commonly found in chromosome analysis at prenatal diagnosis and these are generally considered 'harmless'. It is not clear whether these tetraploid cells simply tend to arise during in vitro cell culture or whether they are also present in placental cells in vivo. There are, at any rate, very few clinical reports of fetuses/infants diagnosed with tetraploidy mosaicism.
Mixoploidy is quite commonly observed in human preimplantation embryos and includes haploid/diploid as well as diploid/tetraploid mixed cell populations. It is unknown whether these embryos fail to implant and are therefore rarely detected in ongoing pregnancies or if there is simply a selective process favoring the diploid cells.
Polyploidy is pervasive in plants and some estimates suggest that 30–80% of living plant species are polyploid, and many lineages show evidence of ancient polyploidy (paleopolyploidy) in their genomes.[21][22][23] Huge explosions in angiosperm species diversity appear to have coincided with the timing of ancient genome duplications shared by many species.[24] It has been established that 15% of angiosperm and 31% of fern speciation events are accompanied by ploidy increase.[25]
Polyploid plants can arise spontaneously in nature by several mechanisms, including meiotic or mitotic failures, and fusion of unreduced (2n) gametes.[26] Both autopolyploids (e.g. potato [27]) and allopolyploids (e.g. canola, wheat, cotton) can be found among both wild and domesticated plant species.
Most polyploids display novel variation or morphologies relative to their parental species, that may contribute to the processes of speciation and eco-niche exploitation.[22][26] The mechanisms leading to novel variation in newly formed allopolyploids may include gene dosage effects (resulting from more numerous copies of genome content), the reunion of divergent gene regulatory hierarchies, chromosomal rearrangements, and epigenetic remodeling, all of which affect gene content and/or expression levels.[28][29][30][31] Many of these rapid changes may contribute to reproductive isolation and speciation. However seed generated from interploidy crosses, such as between polyploids and their parent species, usually suffer from aberrant endosperm development which impairs their viability,[32][33] thus contributing to polyploid speciation.
Lomatia tasmanica is an extremely rare Tasmanian shrub that is triploid and sterile; reproduction is entirely vegetative, with all plants having the same genetic constitution.
There are few naturally occurring polyploid conifers. One example is the Coast Redwood Sequoia sempervirens, which is a hexaploid (6x) with 66 chromosomes (2n = 6x = 66), although the origin is unclear.[34]
Aquatic plants, especially the Monocotyledons, include a large number of polyploids.[35]
The induction of polyploidy is a common technique to overcome the sterility of a hybrid species during plant breeding. For example, Triticale is the hybrid of wheat (Triticum turgidum) and rye (Secale cereale). It combines sought-after characteristics of the parents, but the initial hybrids are sterile. After polyploidization, the hybrid becomes fertile and can thus be further propagated to become triticale.
In some situations, polyploid crops are preferred because they are sterile. For example, many seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual techniques, such as grafting.
Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine.
- Triploid crops: apple, banana, citrus, ginger, watermelon[36]
- Tetraploid crops: apple, durum or macaroni wheat, cotton, potato, canola/rapeseed, leek, tobacco, peanut, kinnow, Pelargonium
- Hexaploid crops: chrysanthemum, bread wheat, triticale, oat, kiwifruit[7]
- Octaploid crops: strawberry, dahlia, pansies, sugar cane, oca (Oxalis tuberosa)[37]
- Dodecaploid crops: some sugar cane hybrids [38]
Some crops are found in a variety of ploidies: tulips and lilies are commonly found as both diploid and triploid; daylilies (Hemerocallis cultivars) are available as either diploid or tetraploid; apples and kinnows can be diploid, triploid, or tetraploid.
Besides plants and animals, the evolutionary history of various fungal species is dotted by past and recent whole-genome duplication events (see Albertin and Marullo 2012[39] for review). Several examples of polyploids are known:
- autopolyploid: the aquatic fungi of genus Allomyces,[40] some Saccharomyces cerevisiae strains used in bakery,[41] etc.
- allopolyploid: the widespread Cyathus stercoreus,[42] the allotetraploid lager yeast Saccharomyces pastorianus,[43] the allotriploid wine spoilage yeast Dekkera bruxellensis,[44] etc.
- paleopolyploid: the human pathogen Rhizopus oryzae,[45] the Saccharomyces genus,[46] etc.
In addition, polyploidy is frequently associated with hybridization and reticulate evolution that appear to be highly prevalent in several fungal taxa. Indeed, homoploid speciation (i.e., hybrid speciation without a change in chromosome number) has been evidenced for some fungal species (e.g., the basidiomycota Microbotryum violaceum [47]).
As for plants and animals, fungal hybrids and polyploids display structural and functional modifications compared to their progenitors and diploid counterparts. In particular, the structural and functional outcomes of polyploid Saccharomyces genomes strikingly reflect the evolutionary fate of plant polyploid ones. Large chromosomal rearrangements[48] leading to chimeric chromosomes[49] have been described, as well as more punctual genetic modifications such as gene loss.[50] The homoealleles of the allotetraploid yeast S. pastorianus show unequal contribution to the transcriptome.[51] Phenotypic diversification is also observed following polyploidization and/or hybridization in fungi,[52] producing the fuel for natural selection and subsequent adaptation and speciation.
Other eukaryotic taxa have experienced one or more polyploidization events during their evolutionary history (see Albertin and Marullo, 2012[39] for review). The oomycetes, which are non-true fungi members, contain several examples of paleopolyploid and polyploid species, such as within the Phytophthora genus.[53] Some species of brown algae (Fucales, Laminariales [54] and diatoms [55]) contain apparent polyploid genomes. In the Alveolata group, the remarkable species Paramecium tetraurelia underwent three successive rounds of whole-genome duplication [56] and established itself as a major model for paleopolyploid studies.
Autopolyploids are polyploids with multiple chromosome sets derived from a single species. Autopolyploids can arise from a spontaneous, naturally occurring genome doubling, like the potato.[27] Others might form following fusion of 2n gametes (unreduced gametes). Bananas and apples can be found as autotriploids. Autopolyploid plants typically display polysomic inheritance, and therefore have low fertility, but may be propagated clonally.
Allopolyploids are polyploids with chromosomes derived from different species. Precisely it is the result of multiplying the chromosome number in an F1 hybrid. Triticale is an example of an allopolyploid, having six chromosome sets, allohexaploid, four from wheat (Triticum turgidum) and two from rye (Secale cereale). Amphidiploids are a type of allopolyploids (they are tetraploid, containing the diploid chromosome sets of both parents[57]). Some of the best examples of allopolyploids come from the Brassicas, and the Triangle of U describes the relationships between the three common diploid Brassicas (B. oleracea, B. rapa, and B. nigra) and three allotetraploids (B. napus, B. juncea, and B. carinata) derived from hybridization among the diploids.
Ancient genome duplications probably occurred in the evolutionary history of all life. Duplication events that occurred long ago in the history of various evolutionary lineages can be difficult to detect because of subsequent diploidization (such that a polyploid starts to behave cytogenetically as a diploid over time) as mutations and gene translations gradually make one copy of each chromosome unlike the other copy. Over time, it is also common for duplicated copies of genes to accumulate mutations and become inactive pseudogenes.[58]
In many cases, these events can be inferred only through comparing sequenced genomes. Examples of unexpected but recently confirmed ancient genome duplications include baker's yeast (Saccharomyces cerevisiae), mustard weed/thale cress (Arabidopsis thaliana), rice (Oryza sativa), and an early evolutionary ancestor of the vertebrates (which includes the human lineage) and another near the origin of the teleost fishes. Angiosperms (flowering plants) have paleopolyploidy in their ancestry. All eukaryotes probably have experienced a polyploidy event at some point in their evolutionary history.
A karyotype is the characteristic chromosome complement of a eukaryote species.[59][60] The preparation and study of karyotypes is part of cytology and, more specifically, cytogenetics.
Although the replication and transcription of DNA is highly standardized in eukaryotes, the same cannot be said for their karotypes, which are highly variable between species in chromosome number and in detailed organization despite being constructed out of the same macromolecules. In some cases, there is even significant variation within species. This variation provides the basis for a range of studies in what might be called evolutionary cytology.
The term is used to describe the relationship between duplicated genes or portions of chromosomes that derived from a common ancestral DNA. Paralogous segments of DNA may arise spontaneously by errors during DNA replication, copy and paste transposons, or whole genome duplications.
The term is used to describe the relationship of similar chromosomes that pair at mitosis and meiosis. In a diploid, one homolog is derived from the male parent (sperm) and one is derived from the female parent (egg). During meiosis and gametogenesis, homologous chromosomes pair and exchange genetic material by recombination, leading to the production of sperm or eggs with chromosome haplotypes containing novel genetic variation.
The term homoeologous, also spelled homeologous, is used to describe the relationship of similar chromosomes or parts of chromosomes brought together following inter-species hybridization and allopolyploidization, and whose relationship was completely homologous in an ancestral species. In allopolyploids, the homologous chromosomes within each parental sub-genome should pair faithfully during meiosis, leading to disomic inheritance; however in some allopolyploids, the homoeologous chromosomes of the parental genomes may be nearly as similar to one another as the homologous chromosomes, leading to tetrasomic inheritance (four chromosomes pairing at meiosis), intergenomic recombination, and reduced fertility.
Example of homoeologous chromosomes[edit]
Durum wheat is the result of the inter-species hybridization of two diploid grass species Triticum urartu and Aegilops speltoides. Both diploid ancestors had two sets of 7 chromosomes, which were similar in terms of size and genes contained on them. Durum wheat contains two sets of chromosomes derived from Triticum urartu and two sets of chromosomes derived from Aegilops speltoides. Each chromosome pair derived from the Triticum urartu parent is homoeologous to the opposite chromosome pair derived from the Aegilops speltoides parent, though each chromosome pair unto itself is homologous.
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- ^ {{cite journal & author = Borneman Anthony R., Zeppel Ryan, Chambers Paul J., Curtin Chris D. & title = Insights into the Dekkera bruxellensis Genomic Landscape: Comparative Genomics Reveals Variations in Ploidy and Nutrient Utilisation Potential amongst Wine Isolates & journal = PLoS Genet & volume = 10 & issue = & pages = e1004161 & year = 2014 & month = & pmid = & doi = 10.1371/journal.pgen.1004161 & url = & isnn = }}
- ^ Ma, Li-Jun; Ibrahim, Ashraf S.; Skory, Christopher; Grabherr, Manfred G.; Burger, Gertraud; Butler, Margi; Elias, Marek; Idnurm, Alexander; Lang, B. Franz; Sone, Teruo; Abe, Ayumi; Calvo, Sarah E.; Corrochano, Luis M.; Engels, Reinhard; Fu, Jianmin; Hansberg, Wilhelm; Kim, Jung-Mi; Kodira, Chinnappa D.; Koehrsen, Michael J.; Liu, Bo; Miranda-Saavedra, Diego; O'Leary, Sinead; Ortiz-Castellanos, Lucila; Poulter, Russell; Rodriguez-Romero, Julio; Ruiz-Herrera, José; Shen, Yao-Qing; Zeng, Qiandong; Galagan, James; Birren, Bruce W. (2009). Madhani, Hiten D, ed. "Genomic Analysis of the Basal Lineage Fungus Rhizopus oryzae Reveals a Whole-Genome Duplication". PLoS Genetics 5 (7): e1000549. doi:10.1371/journal.pgen.1000549. PMC 2699053. PMID 19578406.
- ^ Wong, S.; Butler, G.; Wolfe, K. H. (2002). "Gene order evolution and paleopolyploidy in hemiascomycete yeasts". Proceedings of the National Academy of Sciences 99 (14): 9272–7. Bibcode:2002PNAS...99.9272W. doi:10.1073/pnas.142101099. JSTOR 3059188. PMC 123130. PMID 12093907.
- ^ Devier, B.; Aguileta, G.; Hood, M. E.; Giraud, T. (2009). "Using phylogenies of pheromone receptor genes in the Microbotryum violaceum species complex to investigate possible speciation by hybridization". Mycologia 102 (3): 689–96. doi:10.3852/09-192. PMID 20524600.
- ^ Dunn, B.; Sherlock, G. (2008). "Reconstruction of the genome origins and evolution of the hybrid lager yeast Saccharomyces pastorianus". Genome Research 18 (10): 1610–23. doi:10.1101/gr.076075.108. PMC 2556262. PMID 18787083.
- ^ Nakao, Y.; Kanamori, T.; Itoh, T.; Kodama, Y.; Rainieri, S.; Nakamura, N.; Shimonaga, T.; Hattori, M.; Ashikari, T. (2009). "Genome Sequence of the Lager Brewing Yeast, an Interspecies Hybrid". DNA Research 16 (2): 115–29. doi:10.1093/dnares/dsp003. PMC 2673734. PMID 19261625.
- ^ Scannell, Devin R.; Byrne, Kevin P.; Gordon, Jonathan L.; Wong, Simon; Wolfe, Kenneth H. (2006). "Multiple rounds of speciation associated with reciprocal gene loss in polyploid yeasts". Nature 440 (7082): 341–5. doi:10.1038/nature04562. PMID 16541074.
- ^ Minato, Toshiko; Yoshida, Satoshi; Ishiguro, Tatsuji; Shimada, Emiko; Mizutani, Satoru; Kobayashi, Osamu; Yoshimoto, Hiroyuki (2009). "Expression profiling of the bottom fermenting yeastSaccharomyces pastorianusorthologous genes using oligonucleotide microarrays". Yeast 26 (3): 147–65. doi:10.1002/yea.1654. PMID 19243081.
- ^ Lidzbarsky, Gabriel A.; Shkolnik, Tamar; Nevo, Eviatar (2009). Idnurm, Alexander, ed. "Adaptive Response to DNA-Damaging Agents in Natural Saccharomyces cerevisiae Populations from "Evolution Canyon", Mt. Carmel, Israel". PLoS ONE 4 (6): e5914. doi:10.1371/journal.pone.0005914. PMC 2690839. PMID 19526052.
- ^ Ioos, Renaud; Andrieux, Axelle; Marçais, Benoît; Frey, Pascal (2006). "Genetic characterization of the natural hybrid species Phytophthora alni as inferred from nuclear and mitochondrial DNA analyses". Fungal Genetics and Biology 43 (7): 511–29. doi:10.1016/j.fgb.2006.02.006. PMID 16626980.
- ^ Phillips, N.; Kapraun, D. F.; Gómez Garreta, A.; Ribera Siguan, M. A.; Rull, J. L.; Salvador Soler, N.; Lewis, R.; Kawai, H. (2011). "Estimates of nuclear DNA content in 98 species of brown algae (Phaeophyta)". AoB Plants 2011: plr001. doi:10.1093/aobpla/plr001. PMC 3064507. PMID 22476472.
- ^ Chepurnov, Victor A.; Mann, David G.; Vyverman, Wim; Sabbe, Koen; Danielidis, Daniel B. (2002). "Sexual Reproduction, Mating System, and Protoplast Dynamics of Seminavis (Bacillariophyceae)". Journal of Phycology 38 (5): 1004–19. doi:10.1046/j.1529-8817.2002.t01-1-01233.x.
- ^ Aury, Jean-Marc; Jaillon, Olivier; Duret, Laurent; Noel, Benjamin; Jubin, Claire; Porcel, Betina M.; Ségurens, Béatrice; Daubin, Vincent; Anthouard, Véronique; Aiach, Nathalie; Arnaiz, Olivier; Billaut, Alain; Beisson, Janine; Blanc, Isabelle; Bouhouche, Khaled; Câmara, Francisco; Duharcourt, Sandra; Guigo, Roderic; Gogendeau, Delphine; Katinka, Michael; Keller, Anne-Marie; Kissmehl, Roland; Klotz, Catherine; Koll, France; Le Mouël, Anne; Lepère, Gersende; Malinsky, Sophie; Nowacki, Mariusz; Nowak, Jacek K.; Plattner, Helmut (2006). "Global trends of whole-genome duplications revealed by the ciliate Paramecium tetraurelia". Nature 444 (7116): 171–8. doi:10.1038/nature05230. PMID 17086204.
- ^ Rieger, R.; Michaelis, A.; Green, M.M. (1968). A glossary of genetics and cytogenetics: Classical and molecular.[page needed]
- ^ Edger, Patrick P.; Pires, Chris J. (2009). "Gene and genome duplications: the impact of dosage-sensitivity on the fate of nuclear genes". Chromosome Research 17 (5): 699–717. doi:10.1007/s10577-009-9055-9. PMID 19802709.
- ^ White M.J.D. 1973. The chromosomes. 6th ed, Chapman & Hall, London. p28
- ^ Stebbins G.L. 1950. Variation and evolution in plants. Chapter XII: The Karyotype. Columbia University Press N.Y.[page needed]
Further reading[edit]
- Snustad, D. Peter et al. (2006). Principles of Genetics (4th ed.). Hoboken, NJ: John Wiley & Sons. ISBN 0-471-69939-X.
- The Arabidopsis Genome Initiative (2000). "Analysis of the genome sequence of the flowering plant Arabidopsis thaliana". Nature 408 (6814): 796–815. doi:10.1038/35048692. PMID 11130711.
- Eakin, Guy S.; Behringer, Richard R. (2003). "Tetraploid development in the mouse". Developmental Dynamics 228 (4): 751–66. doi:10.1002/dvdy.10363. PMID 14648853.
- Gaeta, R. T.; Pires, J. C.; Iniguez-Luy, F.; Leon, E.; Osborn, T. C. (2007). "Genomic Changes in Resynthesized Brassica napus and Their Effect on Gene Expression and Phenotype". The Plant Cell Online 19 (11): 3403. doi:10.1105/tpc.107.054346.
- Gregory, T.R.; Mable, B.K. (2005). "Polyploidy in animals". In Gregory, T.R. The Evolution of the Genome. San Diego: Elsevier. pp. 427–517.
- Jaillon, Olivier; Aury, Jean-Marc; Brunet, Frédéric; Petit, Jean-Louis et al. (2004). "Genome duplication in the teleost fish Tetraodon nigroviridis reveals the early vertebrate proto-karyotype". Nature 431 (7011): 946–57. doi:10.1038/nature03025. PMID 15496914.
- Paterson, Andrew H.; Bowers, John E.; Van De Peer, Yves; Vandepoele, Klaas (2005). "Ancient duplication of cereal genomes". New Phytologist 165 (3): 658–61. doi:10.1111/j.1469-8137.2005.01347.x. PMID 15720677.
- Raes, Jeroen; Vandepoele, Klaas; Simillion, Cedric; Saeys, Yvan; Van De Peer, Yves (2003). "Investigating ancient duplication events in the Arabidopsis genome". Journal of Structural and Functional Genomics 3 (1–4): 117–29. doi:10.1023/A:1022666020026. PMID 12836691.
- Simillion, C.; Vandepoele, K; Van Montagu, MC; Zabeau, M; Van De Peer, Y (2002). "The hidden duplication past of Arabidopsis thaliana". Proceedings of the National Academy of Sciences 99 (21): 13627–32. Bibcode:2002PNAS...9913627S. doi:10.1073/pnas.212522399. JSTOR 3073458. PMC 129725. PMID 12374856.
- Soltis, Douglas E.; Soltis, Pamela S.; Schemske, Douglas W.; Hancock, James F.; Thompson, John N.; Husband, Brian C.; Judd, Walter S. (2007). "Autopolyploidy in Angiosperms: Have We Grossly Underestimated the Number of Species?". Taxon 56 (1): 13–30. JSTOR 25065732.
- Soltis DE, Buggs RJA, Doyle JJ, Soltis PS (2010). "What we still don't know about polyploidy". Taxon 59: 1387–403.
- Taylor, J. S.; Braasch, I; Frickey, T; Meyer, A; Van De Peer, Y (2003). "Genome Duplication, a Trait Shared by 22,000 Species of Ray-Finned Fish". Genome Research 13 (3): 382–90. doi:10.1101/gr.640303. PMC 430266. PMID 12618368.
- Tate, J.A.; Soltis, D.E.; Soltis, P.S. (2005). "Polyploidy in plants". In Gregory, T.R. The Evolution of the Genome. San Diego: Elsevier. pp. 371–426.
- Van De Peer, Yves; Taylor, John S.; Meyer, Axel (2003). "Are all fishes ancient polyploids?". Journal of Structural and Functional Genomics 3 (1–4): 65–73. doi:10.1023/A:1022652814749. PMID 12836686.
- Van De Peer, Yves (2004). "Tetraodon genome confirms Takifugu findings: Most fish are ancient polyploids". Genome Biology 5 (12): 250. doi:10.1186/gb-2004-5-12-250. PMC 545788. PMID 15575976.
- Van de Peer, Y.; Meyer, A. (2005). "Large-scale gene and ancient genome duplications". In Gregory, T.R. The Evolution of the Genome. San Diego: Elsevier. pp. 329–68.
- Wolfe, Kenneth H.; Shields, Denis C. (1997). "Molecular evidence for an ancient duplication of the entire yeast genome". Nature 387 (6634): 708–13. doi:10.1038/42711. PMID 9192896.
- Wolfe, Kenneth H. (2001). "Yesterday's polyploids and the mystery of diploidization". Nature Reviews Genetics 2 (5): 333–41. doi:10.1038/35072009. PMID 11331899.
External links[edit]
- Polyploidy on Kimball's Biology Pages
- The polyploidy portal a community-editable project with information, research, education, and a bibliography about polyploidy.


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Considering a Visit?
Alpha Baptist Church is a church in Morristown, Tennessee, that is working to reach our community, nation, and world with the Gospel of Jesus Christ. We would love to have you join us this Sunday or Wednesday and to be a part of all that God is doing in our church. Click here for a virtual tour of some of our facilities.Thinking About Coming? We would love to have you!
You might be wondering...
What Do I Wear? Please come to worship wearing whatever makes you and your family comfortable. Some families choose to wear dress clothes, and just as many dress casually for Sunday morning, Sunday evening, and Wednesday evening services.
What Kind of Music Can I Expect? We use all styles of music to worship Jesus at Alpha. We sing and play traditional hymns, praise and worship choruses, Southern Gospel, and contemporary music in our worship services. Scott Foshie, our Minister of Worship, along with our Celebration Choir and Orchestra, lead us in worship on Sunday mornings and Sunday evenings.
What is the Preaching Like? Pastor Terry Kirby provides Bible teaching from God's Word in all of our worship services. You can listen to some of Pastor Terry's messages in our sermon archive by clicking here.
What is the Best Way to Get to Know People? Sunday School is the heart of our growth at Alpha Baptist Church. We have many different classes for all ages. Sunday School is where you can make new friendships, as well as grow closer to Jesus through Bible study. Sunday School meets at 9 AM every Sunday morning. Please send us an e-mail or give us a call at 423-581-3237 if you'd like to know more about it.
Do You Provide Childcare? We love to minister to your whole family! Loving childcare is provided in the nursery, located close to the sanctuary.
What Ministries Do You Have for My Family? We have many ministries for the entire family. Please click here to see our weekly schedule, or you can call the church office at 423-581-3237, or e-mail us to find out more.
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Celebration Choir Rehearsal for Adults and Middle School: 7:15 - 8:30 PM (Sanctuary)
somethinG BeyOnD College Ministry: 9:00 PM (The Summit)
If you have any questions about our Service Schedule, please call the church office at 423-581-3237 or e-mail us at email@example.com.

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Candid Candace Chicago, hosted by Candace Jordan on Watch312.com, allows viewers to catch a glimpse of what the city's 'who's who' are up to. Ms. Jordan takes you behind the scenes at the awards gala and fundraiser for the Common Ground Foundation, a non-profit that benefits the youth in urban communities.
Common Ground Gala
Posted in: Candid Candace, Videos
– Posted on March 30, 2013


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If there were a Mount Rushmore of movie critics, we'd start with Roger Ebert
BY RICHARD ROEPER SUN-TIMES COLUMNIST April 4, 2013 3:44PM
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Updated: April 8, 2013 4:27PM
Roger would have told me to stop fretting and start writing.
After hearing the terribly sad news of the passing of my writing hero, my friend, my television partner, I sat at the keyboard and tried to come up with the perfect lead to sum up my feelings about Roger Ebert's death.
Five, six minutes went by — an eternity when you're on a self-imposed deadline. And all I could think of was the way Roger's fingers just danced across the keyboard when he was writing a review or filing a story on Oscar night. It was like witnessing a musical genius on a Steinway. His fingers could barely keep up with the narrative flow he was creating on the spot.
Roger was a natural. When it was time to write, he'd sit down and start attacking that keyboard. If there's such a thing as writer's block, I never saw him confronted with it. He was the most prolific, the most gifted, the most dependable newspaperman I've ever known.
If there were a Mount Rushmore of movie critics, we'd start with Roger Ebert, and there would certainly be a place for the late Gene Siskel, and after that there would be room for plenty of debate.
But the discussion would start with Mr. Ebert.
It's impossible to say how many directors, how many writers, how many actors were at least partially inspired to get into the business due to their exposure to "Siskel & Ebert," and the thousands upon thousands of reviews, essays and interviews penned by Roger. All I know is, when I was with Roger at Sundance or the Toronto Film Festival or at the Oscars, it seemed as if not an hour would go by without some well-known film personality approaching to say how much they loved Roger's work, and how much it meant to them and their families when he first mentioned their name in a review.
Shortly after I was named as the successor — I was never the replacement — to Gene Siskel, Roger and I were in New York on a press tour. We were walking through Rockefeller Plaza when a group of Japanese tourists crossed our path.
They immediately stopped, en masse, and started clicking away. Walking around with Roger was like walking around with a live-action cartoon character. People were just thrilled to see him suddenly enter their lives.
He was as famous as the biggest movie stars, as beloved in his home state as the most legendary sports figures. Roger was certainly aware of this, and he was as comfortable with his fame as anyone I've ever met — but it never defined him, it was never something he spent much time pondering.
The guy I spent all those hours with in the screening room, in the TV studio, at film festivals and dinners and traveling, was a guy who loved talking about his beloved wife, Chaz, and his family, a guy who loved telling stories about movie stars and the larger-than-life figures he called his friends, a guy who would wax poetic about his beloved Steak 'n' Shake.
He was political. Everybody knew Roger was a card-carrying liberal, and he was never swayed by the chants of "Stick to movies!" Why should he stick to movies when he was as well-versed and as passionate about politics as half the elected officials in Washington?
He was corny. For years, Roger and Chaz would host massive Fourth of July parties at his home in Michigan, and Roger would always wear his wonderfully tacky American flag shirt while presiding over the karaoke contest and the barbecue and the dancing on the temporary floor installed in the backyard. You never saw him happier than when he was surrounded by family and friends.
He was kind. As a television partner, Roger was exceedingly generous. Even though he was risking the wrath of Disney for spilling the news too soon, Roger told me I had the job before Disney told me I had the job. When the news was made official, Roger took me aside and said, "This is a partnership. You're not a guest on the Roger Ebert show. You're my co-host. It's a 50-50 deal."
And so it was. We had equal time on "Ebert & Roeper." The second time we appeared on "The Tonight Show," Roger insisted it was my turn to take the lead and sit in the chair next to Jay, with Roger on the sofa.
The only thing better than seeing movies with Roger in the screening room on Lake Street in Chicago was talking about movies with Roger in the studio on State Street in Chicago. Years into the job, I'd be sitting there, wondering when someone was going to tap me on the shoulder and tell me to get the hell off the set. To this day, I shake my head in wonder when I look back at all the time I spent with such a great and wonderful presence.
The world lost a great voice today.


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Screening for fraud protects your business from chargebacks and leaves you with more time for real customers. Beagle Lite is a free, customisable tool eWAY offers to all its merchants to block or flag high-risk transactions.
Beagle Verify Allows you to send your customers a customised message via email, SMS or phone alert during checkout to verify their identity. Your customers will need to enter this code during checkout. Beagle Verify is only available on the Responsive Hosted Shared Page.
Beagle ID allows you to customise the information displayed on your customers credit card statement, increasing your brand awareness and increasing customer assurance. This can include your business name, a transaction reference number or your website details. Beagle ID is only available to NAB merchants at this time.
Get the benefits of direct payments without needing to maintain expensive PCI DSS compliance. Client-side encryption auto-encrypts your customer's credit card details through a simple script that is added to your payment form. The customer securely completes the entire transaction on your site - no redirect needed.
Now you can let your customers purchase from your website with a Pre-Coded Pay Now Button. We have already done the work for you - we give you the HTML, and all you need to do is copy and paste it onto your site. Your customers will then be able to click the Pay Now Button with the purchase amount displayed, and pay you using Visa, MasterCard, AMEX, and Diners Club, without leaving your site.
eCommerce is a transaction-intensive industry with high fraud risks. eWAY's Rapid 3.1 API includes a Settlement Report feature that allows merchants to conduct programmatic reconciliation to ensure account figures of purchases and payments are accurate.
Rapid 3.1 transparent redirect allows you to use your own payment page, but transmits card data directly from the customer's browser to eWAY's gateway, bypassing your servers entirely. This transparent redirect model provides enhanced security without impeding your freedom to customise the checkout process.
Your customers want to purchase from you on their mobile devices, so make sure your payment page responds with Responsive Hosted Payments. Create your own hosted payment page that adapts to your customer's preferred device, with beautiful designs and smart templates to boost mobile payment conversions.
Your customers don't have to get their wallets out again with Token Payments. Token is the perfect solution for encouraging return business. Customers can log in to your site and make purchases without having to enter their billing information each time. Token Payments operates through Rapid 3.1 making it fully PCI DSS compliant. For merchants with multiple accounts under one legal business entity, eWAY can generate shared customer tokens across your different websites.
Sell subscriptions or offer payment plans to your customers using Recurring Payments, which automatically charges cards stored within eWAY on a regular basis.
Never leave home without your online store! Turn your mobile device into a virtual point of sale and take payments on the go.
Mobile MYeWAY, a responsive version of the merchant portal, allows you to simply log in, enter customer details, and instantly process transactions. Learn more about Mobile MYeWAY.
For Apple users, eWAY's free mobile app gives you even more options. In addition to taking mobile payments, you can take immediate action on Beagle Alerts and view recent transactions. Download eWAY App from the iTunes Appstore today!
Minimise drop-offs and maximise customer confidence with Direct Payments, which takes customers through the entire transaction without redirecting them away from your website.
Easily take payments via fax, over the phone, or even in-store. Just type the credit card details into MYeWAY, and you will receive an instant transaction result.
For additional fraud-protection, you can use 3-D Secure, a tool which requests a PIN from your customers before processing each transaction
Recruit customers all over the world by taking payments in US Dollars, Great British Pounds, Euros, Canadian Dollars, Japanese Yen, New Zealand Dollars, Singapore Dollars and Hong Kong Dollars.
Want to upsell and market more to your customers? Continue your relationship after the sale, use Transactional Emails to build trust and create loyalty. Advertise directly to a captive market and let them engage with you across multiple social networks.
Maximise sales. Social Connect Emails are a great way to share your success with the online world. Your customers can post their purchase on Facebook and Twitter providing mass advertising and sales opportunities for free!
eWAY allows you to partially or completely refund any transaction. There is no time limit, no need to re-enter the customer's information and no waiting.
Depending on your industry, you may need to reserve funds on a customer's card without charging them. eWAY provides PreAuth for this purpose.
Our Query Service API can check the status of any transaction processed within the last 24 hours. This enables you to query transactions through your shopping cart without logging into eWAY.
If you don't want to charge the customer right away, you can use Stored Payments to save their billing information until you're ready to process the payment.
If you have lots of cards to process, you don't have to enter them individually. With Batch Payments, you can upload the card numbers as a CSV or Excel document, and they will all be processed automatically.
Automating phone payments is easy thanks to eWAY's Interactive Voice Response system. Customers can call, enter their credit card details, and the payment will be processed instantly.
By sharing access to your MYeWAY to partners, you can delegate tasks, such as monitoring your transactions or generating reports. Your Accountant, Developer or another partner can request access through their Partner Portal via Mission Control. You can accept requests and manage shared access, through User Security settings in your MYeWAY account.
Get the most from eWAY by meeting an expert
Take credit card payments seamlessly through your website!
Choose a shopping cart that matches your needs.


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When In Rome
Step into Ancient Rome and follow the story of Flavius, a brave and celebrated warrior who has fought myriad ruthless battles and has lived to tell the tale. After being injured badly by an enemy's spear, Flavius is forced to retire his armor and serve his empire not with brawn, but with tenacity and intellect. Help Flavius follow the Emperor's orders to develop his territory, settle the land, build up trades and expand Rome into a flourishing, prosperous empire! Haggle with shopkeepers, collect supplies, clear road obstructions, construct bridges and more in this fantastic time- and resource-management game!
- 41 Challenging Levels
- Organize production and delivery of goods and products
- Build new settlements and towns
Game System Requirements:
- OS: Windows XP/Windows Vista/Windows 7/Windows 8
- CPU: 1.2 GHz
- DirectX: 9.0
- Hard Drive: 191 MB
Game Manager System Requirements:
- Browser: Internet Explorer 7 or later
The Big Fish Guarantee:
- Quality tested and virus free.
- No ads, no adware, no spyware.


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Reaction Innovations Boom-Boom Tubes 9pk - $4.69
Looking for a new flip bait? These tubes are currently taking the California tournament circuits by storm. The Reaction Innovations Dominator Series Boom-Boom Tubes have put Robert Lee in the winner's circle more than once, and they can do the same for you. They are poured with a super-soft, super-salty plastic that will move off of the hook when flippin' to gaurentee a solid hook-up. A lot of guys loose fish when flippin' tubes; however, with the Boom-Boom Tubes from Reaction Innovations you will be swingin' more fish and cashin' more checks. Find a favorite color and go flippin'!
Length | Quantity |
5" | 9 |
Comments: I have been using these tubes for almost 10 years now and they attract fish and big fish. Only issue I have is that Road Kill is no longer available in the 5" version.
From: RM: Brooklyn, NY 4/22/14
Comments: These tubes have won sooo much money out West. Nobody comes close to the softness and scent on a Boom Boom out of the package. The problem? You can never find them. RI makes wonderful stuff, but is terrible at keeping their vendors supplied. I still use my tubes I have left, but most all the good colors are out of stock.
From: Chris: SoCal
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Here's Hoping Some of Hulu's New Shows Are As Good As 'Battleground'
Earlier this week, the New York Times chronicled Hulu's trajectory from an upstart streaming video service into something more in the vein of a traditional TV network with its own original programming, much like its corporate masters. Of course, that change has been in the works since January. But as Hulu and Netflix face increasing difficulty in convincing TV and cable stations to hand-over programming in a timely manner, they've put more of an emphasis on creating its own content to fill that gap. Read More


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Lift Kits
Here in our Bargain Basement Lift Kits section, you can find bargains on all kinds of clearance parts!
If you are looking at an item, get it while you can because it is gone for good once it's out of stock! We are continuously adding new product to this discounted section of our site, so make sure to keep checking back for added deals.
S10/S15 PU 1983-1994 GM 4WD 4" Lift Kit
Regular Price: $1,710.99
Special Price: $1,167.96
Only 1 in stock


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LDProducts.com offers the highest-quality printing supplies available for Okidata printers including laser toner cartridges. We carry printing supplies for all Okidata models including supplies for Oki printers, OKI B Series printers and many more. Browse through our ever-growing inventory by using the printer model lists below.
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Kids' Star Tile Stepping Stone Kitby Milestones
Everything you need to create a celestial stepping stone. This is a perfect afternoon craft--kids can make a fun stone in about 30 minutes.
Kit contains ceramic star and moon and tiles, 3.5 pounds stepping stone mix, glitter, wooden mixing paddle, craft stick for writing, a reusable stepping stone mold, and instructions. Not for children under 3.
Ask a question about Kids' Star Tile Stepping Stone Kit Ask a question about this product Be the first to write a review on this product! Please wait...
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Processing Time - Orders typically ship in 2-5 business days. Transit via UPS Ground can take up to 6 business days. Braided and custom-made products may take 2 to 3 weeks for delivery.
Free Shipping to Contiguous United States - We do not ship to Alaska, Hawaii or the Caribbean Islands at this time.
Oversized items - Orders that ship freight may take up to 2 weeks from shipment date to be delivered. You will be contacted by the trucking company to schedule a delivery appointment.
Shipping to Canada - Orders shipping to Canada will be subject to a 15% shipping surcharge fee. The shipping surcharge does not cover duties, customs, taxes or brokerage fees which may be charged to you by the shipping courier at time of delivery. At this time we are not shipping to Yukon and the Northwest Territories in Canada.
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Bashian Greenwich Fantasy Ivory Rug
Item #: 362R129IVHG250-P Add to Wishlist
Description: Contemporary patterns in updated colors, highlighted by shimmering viscose, gives this tufted collection a stunning appearance at an incredible value.
- Collection: Greenwich
- Brand: Bashian
- Material: Wool, Viscose
- Weave: Hand Tufted
- Styles: Contemporary Rugs
- Origin: India
- Available in the following sizes: 2' 6" x 8', 3' 9" x 5' 9", 5' 6" x 8' 6", 7' 9" x 9' 9" and 8' 6" x 11' 6"
This is a beautiful rug (runner) and Excellent Quality; HOWEVER, it is too thick and even with a Pad underneath, it is likely to cause one to trip. It is this reason only that causes me to return the rug and give it an average rating.
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RECOMMENDED: Yes, I would recommend this to a friend.This is a good rug. Looks exactly similar to the picture. Sheds a lot and i expected that. Good thickness and smooth. We were worried about the feel of the rug as all of them look great and its hard to buy online. But this is a good buy
Fill out the information below.


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Funeral services will be held at 2:30 p.m. on Saturday, January 5, 2012, at Pilgrim Rest Church in Shelby. Burial will follow at Pilgrim Rest Cemetery in Winstonville with Reverend Spencer Gunn officiating.
Visitation will be from 2-6 p.m. on Friday, January 4, 2013, at Byas Funeral Home in Shelby.
He is survived by his mother, Lillie B. Sutton of Shelby; wife, Gloria Baker of Shelby; daughters, Antrenice Hall, Cassandra Smith, Davetta Powell, and Doneshia Baker of Shelby; brothers, Lendell James of Bartlett, Tenn., Lester Bell of Chicago, Illinois, Michael Morton of Country Club, Illinois, and Jefferson and Alfred Smith of Decatur, Illinois; sisters, Miriam Brown of Merigold, Terbia Daniels of Renova, Lucreatis Bradley of North Ridgeland Hill, Texas, Myrtle and Tara James of Mound Bayou, Helen Morton of Havelcrest, Illinois, and Sandra Jones of Beulah; five grandchildren; and a host of other relatives and friends.


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UTAS Home › Faculty of Arts › School of Humanities › Home › Philosophy & Gender Studies
Philosophy at UTAS is decidedly pluralistic in its outlook. We highly value interdisciplinary collaboration, internationally, nationally, and locally, and we are proud of our friendly and student-centered culture. The discipline of Philosophy contains a number of different traditions, and they are well represented at UTAS. Our program is the only one in Australia that is active in the traditions of Analytic Philosophy, Continental Philosophy, Buddhist Philosophy, Gender Studies, and History of Philosophy.
Our highly ranked teaching and research covers a wide range of topics, including:
Authorised by the Acting Head of School, Humanities
15 January, 2014
Future Students | International Students | Postgraduate Students | Current Students
© University of Tasmania, Australia ABN 30 764 374 782 CRICOS Provider Code 00586B
Copyright | Privacy | Disclaimer | Web Accessibility | Site Feedback | Info line 1300 363 864


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Deleted Scenes
Fans July 24th, 2009
To steal a page from Deadspin here are some deleted scenes, stuff not good enough to get their own post.
- One of the first "big" posts we ever had was this tribute to Bob Uecker. We were linked on Deadspin and a lot of people found us for the first time thanks to that post. Originally there was a line in there that said "Bob has a stalker. Somehow I don't see anyone stalking Ken Harrelson.", but this was later changed. Why? This email:
To: The Editor of Millerparkdrunk.com
Your website posting here:
Created on February 12, 2009 is posting an expunged record associated with my name as part of a listing. I am asking out of professional courtesy that you please remove it from the article. The allegations that you reference in item ten were false, anyway, and the posting you make can be edited to use something else in item 10. Read the rest of this entry »


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CPS wants pre-Labor Day start for school year
BY ROSALIND ROSSI Staff firstname.lastname@example.com January 18, 2013 1:14PM
Chicago Public Schools CEO Barbara Byrd-Bennett speaking at the Chicago Urban League 4510 S. Michigan Friday, November 2, 2012. I Brian Jackson~Sun-Times
Related Documents
Related Stories
Updated: February 20, 2013 6:11AM
Chicago public schools will start classes on Aug. 26 this coming school year, ending a long-held practice of a post-Labor Day opening, under a proposal announced Friday.
Classes will end on June 10; spring break will span April 14-18, and half-days for teacher training will be scrapped.
That's the result of months of discussion following a mutual agreement by CPS officials and the Chicago Teachers Union to dump the current "year-round'' school calendar that begins in August and the "traditional" calendar, that starts after Labor Day, in favor of one "unified" calendar.
School board members must still approve the proposal.
"We are one district, with one vision, and we want to provide our school community with one calendar that makes sense from the classroom to the kitchen table,'' Schools CEO Barbara Byrd-Bennett said of the proposal in a news release.
"Families all across our city struggle to coordinate busy schedules with school, work and play and this unified calendar was designed to best meet the needs of our families' home lives while giving our schools the time they need to implement a rigorous curriculum."
The new calendar also would provide for a five-day Thanksgiving holiday, starting Wed., Nov. 20, and a winter break that would run from Dec. 23 to Jan. 3. Report card pickup would occur on the same day — rather than separate days — for elementary and high school students.
The proposal follows a CPS online survey of more than 13,000 parents, students, teachers and principals in a system that serves 403,000 students.
Teachers who answered the survey overwhelmingly supported a school calendar with an August start — 80 percent of them favored it, according to a CPS summary of the survey. A far smaller 51 percent of surveyed parents and students favored an August start, CPS officials said.
In addition, various calendar options were discussed with teachers union officials and focus groups of parents and principals, CPS said.


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Arizona women's golf added two new players, Gigi Stoll and Danielle Lee, to its roster for the 2015-16 season during this week's National Letter of Intent signing period.
Arizona women's golf will compete at the Pac-12 Preview in Hawaii.
Arizona women's golf completed the final round at the Landfall Tradition.
Arizona women's golf completed day two at the Landfall Tradition.
Arizona women's golf completes the first round at the Landfall Tradition.
Arizona women's golf will travel to North Carolina for the Landfall Tradition.
Arizona will play next at the Landfall Tradition, hosted by UNCW, October 24-26.
Round three at the Windy City Collegiate Classic starts at 5:30 a.m. PT/7:30 a.m. CT on Tuesday.
For the second-straight tournament, Arizona's lineup will include Manon Gidali, Kendall Prince, Krystal Quihuis, Lindsey Weaver and Wanasa Zhou.
The Wildcats shot an 861 (-3) over three rounds in Orlando, Fla.


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2014 Special Events ticket pricing is subject to change and items are subject to availability. We encourage you to buy early and lock in the special item and discounted price.
All Special Event tickets will have a special code on the ticket and are in a specific seating location. In order to redeem any item or gain access to any special activity you will need to present us with your Special Event ticket at the distribution location specified on the Special Events webpage.
How do I know I purchased the correct ticket?
How do I receive my special event item?
Am I guaranteed to receive the special event giveaway item?
I already have tickets to the special event game; can I exchange them for special event tickets?
Can I select a Special Event in my Rays Flex Pack?
Can I purchase the special event item at the game?
If I can't make the game am I able to collect my special event item?
Is there a limit to how many special event tickets I can buy?
If you have any other questions, please call Carmela Wassmer at 727-825-3223 or e-mail firstname.lastname@example.com.


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Despite efforts at the national, European and international level to improve the social and economic integration of Roma in the European Union, many still face deep poverty, profound social exclusion, and discrimination, which often means limited access to quality education, jobs and services, low income levels, sub-standard housing conditions, poor health and lower life expectancy. These problems also present often insurmountable barriers to exercising their fundamental rights.
Data collected systematically over the past years by FRA and international organisations show that the situation of the Roma has changed little in recent years. In the wake of the recent global economic crisis, there is particular concern that the situation may deteriorate further, unless urgent and targeted action is taken.
In April 2011, the European Commission, determined to achieve a tangible change in Roma people's lives over the coming years, published an EU Framework for national Roma integration strategies up to 2020, which set goals at the EU level that require Member States to improve the integration of Roma in four areas:
- employment
- education
- healthcare
- housing.
The Framework called on Member States to submit national Roma integration strategies by the end of 2011, specifying how they will contribute to achieving the overall EU level integration goals, including setting national targets and allowing sufficient funding to deliver them.
On 23 May 2012 the European Commission, having received and assessed national strategies, issued its Communication on National Roma Integration Strategies: a first step in the implementation of the EU Framework. This called on Member States to do more to implement national strategies aimed at improving the economic and social integration of Roma. At the same time, the Commission repeated its request to FRA to continue collecting data across the EU and work closely with Member States to develop effective national monitoring systems.
In response to these developments, FRA developed a multi-annual Roma Programme reaching from 2012 to 2020, to allow the agency to make regular reports on progress made and provide evidence based advice to the EU institutions and Member States based on data systematically collected across the EU. The programme comprises three elements, which are described below.
Collecting data and mapping data sources
- Collecting and analysing primary statistical data through surveys. In 2011, FRA, in close cooperation with the European Commission, the United Nations Development Programme and the World Bank conducted a survey of Roma and non-Roma populations living in close proximity to one another. The study collected data on their socio-economic condition, experiences of discrimination and rights awareness. Further rounds of the survey will be carried out in the future.
- Collecting secondary data and mapping official data sources. Existing data and information are important, as they allow us to contextualise survey findings and provide a more nuanced analysis of the situation. The Agency, through its network of national focal points FRANET, receives and analyses existing official and non-official data and information collected through academic research or by civil society organisations. FRA's research revealed that there are different forms of data collected at various administrative levels, which can be useful in assessing the progress and outcome of Roma integration efforts. The Agency will now map such data sources more systematically, as part of its cooperation with the European Commission and Member States to improve the monitoring of Roma integration efforts.
Collecting data through local engagement
Collecting and analysing primary qualitative data through participatory action research. Starting in 2013, FRA will carry out research at the local level to better understand the factors that influence the current socio-economic situation and life experiences of Roma people, as well as their attitudes, perceptions, values and views. The research will also supply valuable information on how local projects and action plans can best be implemented. The work will entail participatory action research, which means researchers will work in local communities, engaging directly with Roma and non-Roma, and local authorities.
Working with Member States
Assisting Member States to measure progress of Roma integration. From 2012 to 2020, FRA is working closely with the Commission and Member States to develop monitoring methods that can provide a comparative analysis of the situation of Roma across Europe. It is also cooperating with other key actors to assist Member States in developing data collection methodologies, indicators and benchmarks that can enable them to effectively monitor the outcome of policy measures, providing a comparative analysis of the situation of Roma across Europe.
FRA presents Roma Programme progress to national Roma contact points
Widespread Roma exclusion persists, find new surveys
FRA presents its Roma programme
Putting plans to work
Roma Pilot survey
Activity status: Findings available
Start of activity: 2010
In 2011, FRA carried out a survey of Roma in 11 Member States in response to the need to collect comparable data on the socio-economic situation of Roma in the EU. The same information was collected on non-Roma living in areas close by to allow for meaningful comparison.
The EU and international organisations have identified a paucity of data in regard to the socio-economic situation of Roma. FRA research has also shown that Roma are particularly vulnerable to discrimination and human rights violations in the EU. Of all the groups surveyed by FRA in its European Union Minorities and Discrimination Survey (EU-MIDIS), the Roma emerged as the group most likely to face intolerance and social exclusion.
The Roma Pilot Survey, conducted in close collaboration with a parallel survey by the UNDP and the World Bank and funded by the European Commission, produced important data that serve as a baseline for targets to be set by the EU's framework for Roma integration strategies. This survey and subsequent follow-up work is one of the basic pillars of FRA's multi-annual Roma Programme.
The survey applied a similar methodological approach as EU-MIDIS: individuals who identified themselves as Roma were randomly selected and interviewed on the composition of their households, the basic characteristics of household members and their own experiences in key areas related to fundamental rights, for example with respect to social inclusion and integration. Simultaneously, non-Roma living close by were asked the same questions.
Contact: Sabine Springer, Activity manager email@example.com
Collecting secondary data and mapping official data sources
Activity status: Ongoing
Start of activity: 2012
Existing official and non-official data complement and contextualise the statistical results of surveys. In 2012, the Agency asked its FRANET network of national focal points to collect all existing data throughout the Member States related to the socio-economic and human rights situation of Roma.
Data availability and up-to-date information on the current situation of the Roma is a key starting point for identifying goals and targets with which progress can be measured in the future.
In 2013, FRA will map existing data sources at national, regional and selected local levels.
The pilot survey covered only those Member States where Roma live in sufficient numbers and density to allow random sampling. However, FRA needs to analyse the situation of Roma in the remaining 16 Member States and Croatia as well, and will use the existing official and non-official data collected through this process.
In addition, FRA will map existing official and unofficial data sources, including those that may currently be unused or underutilised, in order to support Member States in their efforts to monitor progress made on Roma integration.
FRA collected and analysed the available and relevant data in 2012 [LINK UPCOMING] at national level through FRANET, and is using them in its comparative analyses.
In 2013, FRA will map existing data sources using FRANET and by cooperating with other EU agencies. In addition, FRA will investigate any data produced by research funded by the Socio-economic Sciences and Humanities Programme of the 6th and the 7th Framework Programmes (DG Research).
Contact: Michail Beis, Activity manager email@example.com
Local Engagement for Roma Inclusion (LERI)
Activity status: Ongoing
Start of activity: 2013
The project involves the collection of qualitative data and information through the active involvement of members of local communities – Roma and non-Roma – and local authorities.
This project enables local communities to collect data and shape programmes designed to improve their socio-economic situation and living conditions. It increases our understanding of the situation Roma communities find themselves in and makes it possible to examine and develop ways of improving the design, implementation and monitoring of Roma integration policies and activities at the local level. Statistical data provide essential information, but qualitative in-depth fieldwork research is necessary to better understand the complex interaction of different factors that affect social exclusion processes. The European Commission's Roma Task Force has identified specific weaknesses in the implementation of Roma integration policies and action plans and the use of EU funds, particularly at the local level. The shortcomings included a lack of know-how and administrative capacity at the local level; a lack of reliable data and effective monitoring tools to measure progress; and the lack of meaningful involvement by Roma themselves. The project investigates these deficiencies and the best way of supporting efforts to overcome them.
The research makes use of participatory action research methods. This allows participants to engage directly in the collection and interpretation of data. It also enables those taking part to participate in shaping and monitoring local Roma integration policies and activities. The knowledge and experience gained helps contribute to the capacity of decision makers – particularly at the local level, but also at the national and European levels – to plan and carry out effective and targeted policies that will lead to measurable improvements for Roma.
Contact: Sheena Keller, Activity manager firstname.lastname@example.com
Assisting Member States to measure the progress of Roma integration
Activity status: Ongoing
Start of activity: 2012
FRA has set up and facilitates an ad hoc working party to pool knowledge and expertise on indicator development, data collection, monitoring, and statistical analysis of Roma integration. The working party feeds its results to the European Commission's network of National Roma Contact Points. It is composed of policy and statistical experts from national authorities, the European Commission, EU agencies such as Eurofound, and relevant international bodies such as the UN Development Programme.
Robust monitoring is a key requirement for ensuring a successful implementation of the EU Framework for National Roma Integration. In its Communication of April 2011, the European Commission therefore asked the FRA to work with Member States on developing monitoring methods that can provide a comparative analysis of the situation of Roma throughout the EU.
FRA invited interested EU Member States to participate in a working party that will meet regularly to exchange experience and practices to develop ways of measuring Roma integration so as to produce comparable results. The initial pilot group members are currently Bulgaria, the Czech Republic, Finland, France, Hungary, Italy, The Netherlands, Romania, the Slovak Republic, Spain, the UK and Croatia. This group will be extended over time and lessons learned are regularly provided to all Member States through the Commission's network of National Roma Contact Points.
Contact: Andrey Ivanov, Activity manager firstname.lastname@example.com


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Published on June 7th, 2012 | by Jules IKEAHacker
9Cool teen bedroom workstation
Materials: Expedit, Lack, kitchen cabinets
Description: A young lady transitioning into high school needed a work and entertainment configuration that matched her growing needs. The photo below was how it looked before.
before |
Full specifications and instructions are given at Caylawral's blog.
~ Alison S., Houston, Texas, USA
9 Responses to Cool teen bedroom workstation
Looks great. Nice job!
Please remove the spamcomment above.
spam removed.
Looks so much better
Looks fantastic. I love the set up and the fact that you were super careful in bolting it to the wall. Pinching the idea of cardboard when I next put up new pictures (Rental can't bolt things into the wall even if we could we have a brick house)
Neat work. I am going to bookmark this page for my sons room
You MUST check out her blog post because of all the detail she provides in how the job was done.
Very useful for people not used to thinking in 3D.
Lovely effort.
Wow this looks so good. I want to do that in my bed room on one wall. That would even look good in a kitchen or family room. Really elegant looking. Super wonderful job. I am definitely doing this. You Rock!
How did u get the doors on a bookcase.