Typical structure of a mature eukaryotic mRNA

Polyadenylation is the addition of a poly(A) tail to an RNA molecule. The poly(A) tail consists of multiple adenosine monophosphates; in other words, it is a stretch of RNA that has only adenine bases. In eukaryotes, polyadenylation is part of the process that produces mature messenger RNA (mRNA) for translation. It, therefore, forms part of the larger process of gene expression.

The process of polyadenylation begins as the transcription of a gene finishes, or terminates. The 3'-most segment of the newly-made RNA is first cleaved off by a set of proteins; these proteins then synthesize the poly(A) tail at the RNA's 3' end. In some genes, these proteins may add a poly(A) tail at any one of several possible sites. Therefore, polyadenylation can produce more than one transcript from a single gene (alternative polyadenylation), similar to alternative splicing.^[1]

The poly(A) tail is important for the nuclear export, translation, and stability of mRNA. The tail is shortened over time, and, when it is short enough, the mRNA is enzymatically degraded.^[2] However, in a few cell types, mRNAs with short poly(A) tails are stored for later activation by re-polyadenylation in the cytosol.^[3] In contrast, when polyadenylation occurs in bacteria, it promotes RNA degradation.^[4] This is also sometimes the case for eukaryotic non-coding RNAs.^[5] The wide distribution of polyadenylation among living organisms indicates that this process evolved early in the history of life on Earth.

Background on RNA[link]

Chemical structure of RNA. The sequence of bases differs between RNA molecules.

For further information, see RNA and Messenger RNA

RNAs are a type of large biological molecules, whose individual building blocks are called nucleotides. The name poly(A) tail (for polyadenylic acid tail)^[6] reflects the way RNA nucleotides are abbreviated, with a letter for the base the nucleotide contains (A for adenine, C for cytosine, G for guanine and U for uracil). RNAs are produced (transcribed) from a DNA template. By convention, RNA sequences are written in a 5' to 3' direction. The 5' end is the part of the RNA molecule that is transcribed first, and the 3' end is transcribed last. The 3' end is also where the poly(A) tail is found on polyadenylated RNAs.^[1][7]

Messenger RNA (mRNA) is RNA that has a coding region that acts as a template for protein synthesis (translation). The rest of the mRNA, the untranslated regions, tune how active the mRNA is.^[8] There are also many RNAs that are not translated, called non-coding RNAs. Like the untranslated regions, many of these non-coding RNAs have regulatory roles.^[9]

Nuclear polyadenylation[link]

Function[link]

In nuclear polyadenylation, a poly(A) tail is added to an RNA at the end of transcription. On mRNAs, the poly(A) tail protects the mRNA molecule from enzymatic degradation in the cytoplasm and aids in transcription termination, export of the mRNA from the nucleus, and translation.^[2] Almost all eukaryotic mRNAs are polyadenylated,^[10] with the exception of animal replication-dependent histone mRNAs.^[11] These are the only mRNAs in eukaryotes that lack a poly(A) tail, ending instead in a stem-loop structure followed by a purine-rich sequence, termed histone downstream element, that directs where the RNA is cut so that the 3' end of the histone mRNA is formed.^[12]

Many eukaryotic non-coding RNAs are always polyadenylated at the end of transcription. There are small RNAs where the poly(A) tail is seen only in intermediary forms and not in the mature RNA as the ends are removed during processing, the notable ones being microRNAs.^[13][14] But, for many long noncoding RNAs – a seemingly large group of regulatory RNAs that, for example, includes the RNA Xist, which mediates X chromosome inactivation – a poly(A) tail is part of the mature RNA.^[15]

Mechanism[link]

The polyadenylation machinery in the nucleus of eukaryotes works on products of RNA polymerase II, such as precursor mRNA. Here, a multi-protein complex (see components on the right) cleaves the 3'-most part of a newly produced RNA and polyadenylates the end produced by this cleavage. The cleavage is catalysed by the enzyme CPSF^[11] and occurs 10–30 nucleotides downstream of its binding site.^[16] This site is often the sequence AAUAAA on the RNA, but variants of it that bind more weakly to CPSF exist.^[17] Two other proteins add specificity to the binding to an RNA: CstF and CFI. CstF binds to a GU-rich region further downstream of CPSF's site.^[18] CFI recognises a third site on the RNA (a set of UGUAA sequences in mammals^[19][20][21]) and can recruit CPSF even if the AAUAAA sequence is missing.^[22][23] The polyadenylation signal – the sequence motif recognised by the RNA cleavage complex – varies between groups of eukaryotes. Most human polyadenylation sites contain the AAUAAA sequence,^[18] but this sequence is less common in plants and fungi.^[24]

The RNA is cleaved right after transcription, as CstF also binds to RNA polymerase II.^[25] Cleavage also involves the protein CFII, though it is unknown how.^[26] The cleavage site associated with a polyadenylation signal can vary up to some 50 nucleotides.^[27]

When the RNA is cleaved, polyadenylation starts, catalysed by polyadenylate polymerase. Polyadenylate polymerase builds the poly(A) tail by adding adenosine monophosphate units from adenosine triphosphate to the RNA, cleaving off pyrophosphate.^[28] Another protein, PAB2, binds to the new, short poly(A) tail and increases the affinity of polyadenylate polymerase for the RNA. When the poly(A) tail is approximately 250 nucleotides long the enzyme can no longer bind to CPSF and polyadenylation stops, thus determining the length of the poly(A) tail.^[29][30] CPSF is in contact with RNA polymerase II, allowing it to signal the polymerase to terminate transcription.^[31][32] The polyadenylation machinery is also physically linked to the spliceosome, a complex that removes introns from RNAs.^[23]

Downstream effects[link]

The poly(A) tail acts as the binding site for poly(A)-binding protein. Poly(A)-binding protein promotes export from the nucleus and translation, and inhibits degradation.^[33] This protein binds to the poly(A) tail prior to mRNA export from the nucleus and in yeast also recruits poly(A) nuclease, an enzyme that shortens the poly(A) tail and allows the export of the mRNA. Poly(A)-binding protein is exported to the cytoplasm with the RNA. mRNAs that are not exported are degraded by the exosome.^[34][35] Poly(A)-binding protein also can bind to, and thus recruit, several proteins that affect translation,^[34] one of these is initiation factor-4G, which in turn recruits the 40S ribosomal subunit.^[36] However, a poly(A) tail is not required for the translation of all mRNAs.^[37]

Deadenylation[link]

In eukaryotic somatic cells, the poly(A) tail of most mRNAs in the cytoplasm gradually get shorter, and mRNAs with shorter poly(A) tail are translated less and degraded sooner.^[38] However, it can take many hours before an mRNA is degraded.^[39] This deadenylation and degradation process can be accelerated by microRNAs complementary to the 3' untranslated region of an mRNA.^[40] In immature egg cells, mRNAs with shortened poly(A) tails are not degraded, but are instead stored without being translated. They are then activated by cytoplasmic polyadenylation after fertilisation, during egg activation.^[41]

In animals, poly(A) ribonuclease (PARN) can bind to the 5' cap and remove nucleotides from the poly(A) tail. The level of access to the 5' cap and poly(A) tail is important in controlling how soon the mRNA is degraded. PARN deadenylates less if the RNA is bound by the initiation factors 4E (at the 5' cap) and 4G (at the poly(A) tail), which is why translation reduces deadenylation. The rate of deadenylation may also be regulated by RNA-binding proteins. Once the poly(A) tail is removed, the decapping complex removes the 5' cap, leading to a degradation of the RNA. Several other enzymes that seem to be involved in deadenylation have been identified in yeast.^[42]

Alternative polyadenylation[link]

Results of using different polyadenylation sites on the same gene

Many protein-coding genes have more than one polyadenylation site, so a gene can code for several mRNAs that differ in their 3' end.^[24][43][44] Since alternative polyadenylation changes the length of the 3' untranslated region, it can change which binding sites for microRNAs the 3' untranslated region contains.^[16][45] MicroRNAs tend to repress translation and promote degradation of the mRNAs they bind to, although there are examples of microRNAs that stabilise transcripts.^[46][47] Alternative polyadenylation can also shorten the coding region, thus making the mRNA code for a different protein,^[48][49] but this is much less common than just shortening the 3' untranslated region.^[24]

The choice of poly(A) site can be influenced by extracellular stimuli and depends on the expression of the proteins that take part in polyadenylation.^[50][51] For example, the expression of CstF-64, a subunit of cleavage stimulatory factor (CstF), increases in macrophages in response to lipopolysaccharides (a group of bacterial compounds that trigger an immune response). This results in the selection of weak poly(A) sites and thus shorter transcripts. This removes regulatory elements in the 3' untranslated regions of mRNAs for defense-related products like lysozyme and TNF-α. These mRNAs then have longer half-lives and produce more of these proteins.^[50] RNA-binding proteins other than those in the polyadenylation machinery can also affect whether a polyadenyation site is used,^{[52][53][54][55]}as can DNA methylation near the polyadenylation signal.^[56]

Cytoplasmic polyadenylation[link]

There is polyadenylation in the cytosol of some animal cell types, namely in the germ line, during early embryogenesis and in post-synaptic sites of nerve cells. This lengthens the poly(A) tail of an mRNA with a shortened poly(A) tail, so that the mRNA will be translated.^[38][57] These shortened poly(A) tails are often less than 20 nucleotides, and are lengthened to around 80–150 nucleotides.^[3]

In the early mouse embryo, cytoplasmic polyadenylation of maternal RNAs from the egg cell allows the cell to survive and grow even though transcription does not start until the middle of the 2-cell stage (4-cell stage in human).^[58][59] In the brain, cytoplasmic polyadenylation is active during learning and could play a role in long-term potentiation, which is the strengthening of the signal transmission from a nerve cell to another in response to nerve impulses and is important for learning and memory formation.^[3][60]

Cytoplasmic polyadenylation requires the RNA-binding proteins CPSF and CPEB, and can involve other RNA-binding proteins like Pumilio.^[61] Depending on the cell type, the polymerase can be the same type of polyadenylate polymerase (PAP) that is used in the nuclear process, or the cytoplasmic polymerase GLD-2.^[62]

Tagging for degradation in eukaryotes[link]

For many non-coding RNAs, including tRNA, rRNA, snRNA, and snoRNA, polyadenylation is a way of marking the RNA for degradation, in at least yeast.^[63] This polyadenylation is done in the nucleus by the TRAMP complex, which adds a tail that is around 40 nucleotides long to the 3' end.^[64] The RNA is then degraded by the exosome.^[65] Poly(A) tails have also been found on human rRNA fragments, both the form of homopolymeric (A only) and heterpolymeric (mostly A) tails.^[66]

In prokaryotes and organelles[link]

In many bacteria, both mRNAs and non-coding RNAs can be polyadenylated. This poly(A) tail promotes degradation by the degradosome, which contains two RNA-degrading enzymes: polynucleotide phosphorylase and RNase E. Polynucleotide phosphorylase binds to the 3' end of RNAs and the 3' extension provided by the poly(A) tail allows it to bind to the RNAs whose secondary structure would otherwise block the 3' end. Successive rounds of polyadenylation and degradation of the 3' end by polynucleotide phosphorylase allows the degradosome to overcome these secondary structures. The poly(A) tail can also recruit RNases that cut the RNA in two.^[67] These bacterial poly(A) tails are about 30 nucleotides long.^[68]

In as different groups as animals and trypanosomes, the mitochondria contain both stabilising and destabilising poly(A) tails. Destabilising polyadenylation targets both mRNA and noncoding RNAs. The poly(A) tails are 43 nucleotides long on average. The stabilising ones start at the stop codon, and without them the stop codon (UAA) is not complete as the genome only encodes the U or UA part. Plant mitochondria have only destabilising polyadenylation, and yeast mitochondria have no polyadenylation at all.^[69]

While many bacteria and mitochondria have polyadenylate polymerases, they also have another type of polyadenylation, performed by polynucleotide phosphorylase itself. This enzyme is found in bacteria,^[70] mitochondria,^[71] plastids^[72] and as a constituent of the archeal exosome (in those archaea that have an exosome).^[73] It can synthesise a 3' extension where the vast majority of the bases are adenines. Like in bacteria, polyadenylation by polynucleotide phosphorylase promotes degradation of the RNA in plastids^[74] and likely also archaea.^[69]

Evolution[link]

Although polyadenylation is seen in almost all organisms, it is not universal.^[75][76] However, the wide distribution of this modification and the fact that it is present in organisms from all three domains of life implies that the last universal common ancestor of all living organisms, it is presumed, had some form of polyadenylation system.^[68] A few organisms do not polyadenylate mRNA, which implies that they have lost their polyadenylation machineries during evolution. Although no examples of eukaryotes that lack polyadenylation are known, mRNAs from the bacterium Mycoplasma gallisepticum and the salt-tolerant archaean Haloferax volcanii lack this modification.^[77][78]

The most ancient polyadenylating enzyme is polynucleotide phosphorylase. This enzyme is part of both the bacterial degradosome and the archaeal exosome,^[79] two closely related complexes that recycle RNA into nucleotides. This enzyme degrades RNA by attacking the bond between the 3'-most nucleotides with a phosphate, breaking off a diphosphate nucleotide. This reaction is reversible, and so the enzyme can also extend RNA with more nucleotides. The heteropolymeric tail added by polynucleotide phosphorylase is very rich in adenine. The choice of adenine is most likely the result of higher ADP concentrations than other nucleotides as a result of using ATP as an energy currency, making it more likely to be incorporated in this tail in early lifeforms. It has been suggested that the involvement of adenine-rich tails in RNA degradation prompted the later evolution of polyadenylate polymerases (the enzymes that produce poly(A) tails with no other nucleotides in them).^[80]

Polyadenylate polymerases are not as ancient. They have separately evolved in both bacteria and eukaryotes from CCA-adding enzyme, which is the enzyme that completes the 3' ends of tRNAs. Its catalytic domain is homologous to that of other polymerases.^[65] It is presumed that the horizontal transfer of bacterial CCA-adding enzyme to eukaryotes allowed the archaeal-like CCA-adding enzyme to switch function to a poly(A) polymerase.^[68] Some lineages, like archaea and cyanobacteria, never evolved a polyadenylate polymerase.^[80]

History[link]

Polyadenylation was first identified in 1960 as an enzymatic activity in extracts made from cell nuclei that could polymerise ATP, but not ADP, into polyadenine.^[81][82] Although identified in many types of cells, this activity had no known function until 1971, when poly(A) sequences were found in mRNAs.^[83][84] The only function of these sequences was thought at first to be protection of the 3' end of the RNA from nucleases, but later the specific roles of polyadenylation in nuclear export and translation were identified. The polymerases responsible for polyadenylation were first purified and characterized in the 1960s and 1970s, but the large number of accessory proteins that control this process were discovered only in the early 1990s.^[83]

See also[link]

• Simian virus 40 late polyadenylation signal (SVLPA)

References[link]

Further reading[link]

• Danckwardt, Sven; Hentze, Matthias W; Kulozik, Andreas E (2008). "3′ end mRNA processing: molecular mechanisms and implications for health and disease". The EMBO Journal 27 (3): 482–98. DOI:10.1038/sj.emboj.7601932. PMC 2241648. PMID 18256699. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2241648. 

Poly Styrene
Background information
Birth name	Marianne Joan Elliott-Said
Born	3 July 1957 Bromley, Kent, England
Died	25 April 2011(2011-04-25) (aged 53) Sussex, England
Genres	Punk, rock, New Wave, dub
Occupations	Vocalist, songwriter
Years active	1976–2011
Labels	Universal, EMI/Virgin, Future Noise Music
Associated acts	X-Ray Spex

Poly Styrene was the stage name of Marianne Joan Elliott-Said (3 July 1957 – 25 April 2011),^[1][2] a British musician, songwriter and singer, most notably in the pioneering punk rock band X-Ray Spex.

Early life[link]

Marianne Elliott-Said was born in 1957 in Bromley, Kent, and raised in Brixton, London.^[3] Her mother, who raised her alone, was a Scots-Irish legal secretary.^[4] Her father was a dispossessed Somali aristocrat.^[5][6]

As a teenager, Marianne was a "barefoot hippie". At age 15, she ran away from home with £3 in her pocket, and hitchhiked from one music festival to another, staying at hippie crash pads. She thought of this as a challenge to survive. The adventure ended when she stepped on a rusty nail while bathing in a stream and had to be treated for septicaemia.^[4]

After seeing the Sex Pistols performance at the Pier Pavillion Hastings on 3 July 1976^[7][8] (her nineteenth birthday) she thought that anyone could do what they were doing and so decided to form the punk band X-Ray Spex. She eventually became a passionate feminist and posted a blog dedicated to women's rights and defence of women.

Music career[link]

Poly recorded her first demo with producer Ted Bunting in 1975, when she was just 16 years old,^[9] and released her first, reggae, single, "Silly Billy"/"What A Way", as "Mari Elliott" in 1976.^[10]

After watching a very early gig by the Sex Pistols in an empty hall on Hastings Pier, playing a set of cover songs,^[8] she thought that anyone could do what they were doing and so decided to put an ad in the paper for ‘young punx who want to stick it together’ to form a band.^[11] So it was that, as Poly Styrene, the singer with X-Ray Spex,^[4] she was described by Billboard as the "archetype for the modern-day feminist punk"; because she wore dental braces, stood against the typical sex object female of 1970s rock star, sported a gaudy Dayglo wardrobe, and was of mixed race, she was "one of the least conventional front-persons in rock history, male or female".^[12]

In 1978, after a gig in Doncaster, she had a vision of a pink light in the sky and felt objects crackling when she touched them. Thinking she was hallucinating, her mother took her to the hospital where Marianne was misdiagnosed with schizophrenia, sectioned, and told she would never work again. Although she missed playing at the time, in hindsight, she felt that getting out of the public eye was good for her. She was diagnosed with bipolar disorder in 1991.^[13]

After the original version of X-Ray Spex broke up, Poly Styrene recorded a solo album, Translucence, in 1980. The album abandoned X-Ray Spex's loud guitar work for a quieter and more jazzy sound that has since been described as anticipating later work by Everything but the Girl.^[14] In 1986, she released the EP God's & Godesses [sic] on the Awesome record label. A New Age solo album, Flower Aeroplane, followed in 2004.^[4]

In 1983, she was initiated into the Hare Krishna movement and recorded at their recording studios while living as a devotee at Bhaktivedanta Manor. She lived as a Hare Krishna convert in Hertfordshire and London from 1983 to 1988.^[15]

In 2007, she was invited to the Concrete Jungle festival in Camber Sands,^[16] where she and the gathering's organiser, Symond Lawes, agreed to initiate a 30-year celebration of X-Ray Spex's debut album, Germ Free Adolescents. They decided to hold a live show at the Camden Roundhouse, which was a sell-out event on 6 September 2008. A live album/DVD of this event, Live @ The Roundhouse London 2008, was released in November 2009 on The Year Zero label by Future Noise Music.

She made a guest appearance at the 2008 30th anniversary concert of Rock Against Racism in Victoria Park, London, performing "Oh Bondage Up Yours" with guest musicians Drew McConnell (of Babyshambles and Helsinki) and 'Flash' David Wright playing saxophone.

That same year, she dueted with Goldblade's John Robb on a remix of Goldblade's "City Of Christmas Ghosts".^[17]

In March 2009, she joined other members of PRS for Music in criticising Google for allegedly not paying a fair share of royalties to musicians. This followed Google's removal of millions of videos from YouTube because of a royalties dispute with the organisation.^[18]

NME.com announced on 29 October 2010 that Poly Styrene was to release a solo album titled Generation Indigo, produced by Martin Glover (aka Youth from Killing Joke), in March 2011. She released a free download of "Black Christmas" in November 2010.^[19] "Black Christmas" featured and was written in collaboration with her daughter, Celeste. It was inspired by the killing spree in Los Angeles instigated by a man dressed as Santa Claus.^[20]

Poly Styrene announced "Virtual Boyfriend" as the first single from her new album Generation Indigo via Spinner Music,^[21] as well as the launch of her brand new website.^[22] "Virtual Boyfriend" was released on 21 March 2011, and featured an animated promotional video directed by Ben Wheele. Generation Indigo was released on 28 March 2011, via Future Noise Music. The album received critical acclaim, including a perfect 10 out of 10 score in Artrocker magazine, and 8 out of 10 in The Telegraph newspaper. Generation Indigo was also chosen as Album of the Day on UK radio station BBC 6 Music.

"Ghoulish" was the first posthumous single to be released from Generation Indigo, and was backed by a remix from Hercules and Love Affair.

Personal life[link]

Poly Styrene described herself as "an observer, not a suffering artist writing from tortured experiences. I was playing with words and ideas. Having a laugh about everything, sending it up."^[4]

She lived in St Leonards, East Sussex.^[5]

Her daughter Celeste Bell-Dos Santos is the frontwoman for the music group Debutant Disco based in Madrid, Spain.^[23]

In March 2009, Poly Styrene took part in the inaugural Instigate Debate night. The night's theme was modern day consumerism. Other current issues were also discussed.^[24]

Death[link]

In February 2011, in an interview published in The Sunday Times magazine, which largely focused on her past and present relationship with her daughter Celeste, she revealed that she had been treated for breast cancer, and that it had spread to her spine and lungs. She died on 25 April 2011^[1][2][25] at the age of 53.

Solo discography[link]

Albums[link]

• Translucence (United Artists, 1980)^[26]
• Flower Aeroplane (2004)^[27]
• Generation Indigo (Future Noise Music, 2011)^[28][29]

EPs[link]

• God's & Godesses (Awesome, 1986)^[30]

Singles[link]

• "Silly Billy"/"What A Way" — as Mari Elliot (GTO, 1976)^[10]
• "Talk In Toytown"/"Sub Tropical" (United Artists, 1980)^[31]
• "City Of Christmas Ghosts" — Goldblade featuring Poly Styrene (Damaged Goods, 2008)^[32]
• "Black Christmas" (2010)^[19]
• "Virtual Boyfriend" (2011)^[21]
• "Ghoulish" (2011)

References[link]

External links[link]

• Poly Styrene's official website, Poly Styrene's official web site
• Poly's Punky Party, personal site

Anja Rubik
Anja Rubik walking for Michael Kors Spring/Summer 2009
Born	Anna Rubik (1983-06-12) 12 June 1983 (age 29) Rzeszów[1] Poland
Height	1.79 m (5 ft 10 in)[2]
Hair color	Blonde
Eye color	Blue
Measurements	(US) 33-24-35 ; (EU) 84-61-90[2]
Dress size	(US) 4 ; (EU) 36[2]
Website
http://www.anjarubik.com/

Anja Rubik (born Anna Rubik, 12 June 1983^[1]) is a Polish model.

Early life[link]

Rubik was born in Rzeszów, Poland.^[1] In 1988 she and her family moved to Greece. They later moved to Canada and to Umtata, South Africa. Among the schools she attended was one in Częstochowa, Poland. Rubik was introduced to modeling while attending a British high school in Paris. She wanted to finish her schooling before officially becoming a model, and so only worked on school holidays. Her mother, father and sister still reside in Poland.^[3]

Career[link]

Rubik began her modeling career full-time soon after graduating from high school. She has been featured on the covers of various magazines, including French, Korean, Latin American, Japanese, Spanish, Australian, and German Vogue, Korean and French Numéro, Nylon and Elle. She has modeled for such designers/brands as Shiatzy Chen, Givenchy, Chloé, Christian Dior, Hermès, Valentino, Gucci, H&M, GAP, Lanvin, Balmain, Belstaff, Armani, Lacoste, DKNY, Barneys New York, Jimmy Choo, Estee Lauder, Roberto Cavalli, and Chanel.^[4] She has also done haute couture for Valentino, Chanel, Christian Lacroix and Elie Saab.

Rubik was one of the faces to land in one of the fourteen covers of V magazine September 2008 issue. Each cover boasts a head shot of a famous model, either from the new crop of leading models (Agyness Deyn, Lara Stone, Natasha Poly, Daria Werbowy etc.) or the supermodel era (Christy Turlington, Naomi Campbell, Eva Herzigova), it was lensed by duo Inez van Lamsweerde and Vinoodh Matadin.^[5]

She appeared in the 2009, 2010, and 2011 Victoria's Secret Fashion Shows.

Vogue Paris declared her one of the top 30 models of the 2000s.^[6] As of May 2012^[update] she is ranked 3rd, behind Natasha Poly and Joan Smalls, on the Top 50 Models Women list and 18th on the Top 25 Money Girls list, which ranks the highest earning high fashion models in the industry, by models.com.^[7][8]

Rubik is featured in the 2009 fitness video game Yoga Wii for the Wii home console.^[9] She was also an inspiration for a character in the manga "The One" as the main character Cane Lele.^{[citation needed]}

Since 2010, she has appeared on Top Model. Zostań modelką, the Polish edition of America's Next Top Model.^[10]

Rubik is featured in the 2011 Pirelli Calendar photographed by Karl Lagerfeld. She was one of the models to star in Lacoste's "new look" campaign in January, a different advertising concept for 2011 under the new tagline, “Unconventional Chic”. The ads were shot by Mert and Marcus, showing models wearing the iconic white Lacoste polo shirts worn over fancy black evening wear.^[11] Rubik renewed her contract with Fendi for the third season in a row.^[12]

Personal life[link]

Rubik and her longtime boyfriend, Serbian model Sasha Knezevic, became engaged just before Christmas 2010 in Vienna.^[13] They married on 16 July 2011 in Deia, Majorca.^[14]

References[link]