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Hadrian - Wikipedia

Download date: 22 Nov Latin literature, the recognition of which may shed a new light on its interpretation and text. Fontecedro , Holzhausen , for religious aspects Steinmetz , and Gwyn Griffiths testo latino (Italian Edition) - Kindle edition by Emanuela Andreoni Fontecedro, Publication Date: January 17, ; Sold by: Amazon Digital Services LLC. Guida alla traduzione del testo latino (Italian Edition). 17 Jan | Kindle by Maura Amoroso and Emanuela Andreoni Fontecedro. £Kindle Edition.

We profiled PBOV1 expression in multiple cancer and normal tissue samples and found that it was expressed in 19 out of 34 tumors of various origins but completely lacked expression in any of the normal adult or fetal human tissues.

Our analysis of public microarray data suggests that PBOV1 activation in tumors could be dependent on the Hedgehog signaling pathway. Despite the recent de novo origin and the lack of identifiable functional signatures, a missense SNP in the PBOV1 coding sequence has been previously associated with an increased risk of breast cancer.

Using publicly available microarray datasets, we found that high levels of PBOV1 expression in breast cancer and glioma samples were significantly associated with a positive outcome of the cancer disease.

We also found that PBOV1 was highly expressed in primary but not in recurrent high-grade gliomas, suggesting the presence of a negative selection against PBOV1-expressing cancer cells. Our findings could contribute to the understanding of the mechanisms behind de novo gene origin and the possible role of tumors in this process. Introduction The origin of novel genes in the evolution of multicellular organisms has long been postulated to play a fundamental role in the development of new functions [1].

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There are several well-established mechanisms of novel gene origin. For example, duplication and divergence, retroposition, gene fusion, exon shuffling and horizontal gene transfer all rely on reuse of the pre-existing genetic material see [2] for review. It has been also proposed that some protein-coding genes might have originated de novo from non-coding genomic regions through a series of mutations ultimately leading to the appearance of a novel protein-coding transcript. The resulting proteins might be fixed in the evolution either as a result of genetic drift or due to an accidental positive contribution to the organism fitness.

The positive selection following the fixation might further enhance the functionality of such proteins. Despite the de novo mechanism of gene origin for a long time being considered unrealistic, there is a growing number of reports from various species that show that de novo gene origin is a widespread process that takes place in all branches of the tree of life [3] — [9].

However, the detailed understanding of the de novo gene origin is still missing, including what are the forces that drive the initial fixation of a newly originated gene and how does its function get shaped and integrated into the organism context. We have earlier hypothesized that tumorogenesis may play an important role in the novel gene origin and fixation detailed in [10] and [11].

Briefly, one prominent feature of various tumor types is the abundant upregulation of various transcripts, many of which have an uncharacterized function [12][13]. These genes are controlled by CpG-island based promoters and are activated preferentially in spermatocytes and in various cancer types, whereupon the activation in both cases is linked to a widespread loss of CpG methylation [14][15]. Most of such transcripts lack an established function and are silent in most of the normal tissues.

However, some may happen to have a protein-coding potential and thus can be potentially classified as de novo genes. Second, Fisher and co-workers [16] have showed that some of the proteins from a library of randomly generated protein-coding sequences were able to rescue auxotrophic mutants of E. Although this proof-of-principle example shows that a previously noncoding and non-optimized sequence may readily give rise to a minimally functional protein, we believe that in most cases a recently emerged de novo gene would initially lack functional features such that they would be sufficient to facilitate its evolutionary fixation.

Given the ongoing mutational process and the lack of selective pressure the half-life of such a gene could be relatively short. We hypothesized that the expression of emerging de novo genes in tumors might in some way help to create a phenotypic feedback loop that would facilitate evolutionary fixation of those genes and their further functional integration into the context of the organism.

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With an aim to find specific examples to support this hypothesis, we focused on searching for human evolutionarily novel genes with a preferential expression in tumors. We previously reported several such transcripts, but most of them lacked protein-coding potential [17][18]. The gene has been first characterized by An and co-workers [21] as being overexpressed in prostate, breast, and bladder cancer.

The authors expressed the protein in vitro, produced antibodies and showed that PBOV1 protein was present in the blood of prostate cancer patients but not in the healthy controls. Hadrian's Gatein Antalya, southern Turkey was built to honour Hadrian who visited the city in Arch of Hadrian in JerashTransjordanbuilt to honour Hadrian's visit in Prior to Hadrian's arrival in Britanniathe province had suffered a major rebellion, from to Fronto writes about military losses in Britannia at the time.

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In Hadrian initiated the construction of a wall, "to separate Romans from barbarians". Reduction of defense costs may also have played a role, as the Wall deterred attacks on Roman territory at a lower cost than a massed border army, [79] and controlled cross-border trade and immigration.

He never saw the finished wall that bears his name. Hadrian appears to have continued through southern Gaul. At Nemausushe may have overseen the building of a basilica dedicated to his patroness Plotina, who had recently died in Rome and had been deified at Hadrian's request. At some point, he visited Cyrenewhere he personally funded the training of young men from well-bred families for the Roman military.

Cyrene had benefited earlier in from his restoration of public buildings destroyed during the earlier Jewish revolt. Nicomedia had been hit by an earthquake only shortly before his stay; Hadrian provided funds for its rebuilding, and was acclaimed as restorer of the province. Literary and epigraphic sources say nothing on when or where they met; depictions of Antinous show him aged 20 or so, shortly before his death in In he would most likely have been a youth of 13 or Various traditions suggest his presence at particular locations, and allege his foundation of a city within Mysia, Hadrianutheraeafter a successful boar hunt.

At about this time, plans to complete the Temple of Zeus in Cyzicusbegun by the kings of Pergamonwere put into practice. The temple received a colossal statue of Hadrian. Hadrian arrived in Greece during the autumn ofand participated in the Eleusinian Mysteries. He refused to intervene in a local dispute between producers of olive oil and the Athenian Assembly and Councilwho had imposed production quotas on oil producers; [96] yet he granted an imperial subsidy for the Athenian grain supply.

Antinous and Hadrian may have already been lovers at this time; Hadrian showed particular generosity to Mantineawhich shared ancient, mythic, politically useful links with Antinous' home at Bithynia.

He restored Mantinea's Temple of Poseidon Hippios[] [] and according to Pausanias, restored the city's original, classical name. Hadrian also rebuilt the ancient shrines of Abae and Megaraand the Heraion of Argos. The two aristocrats would be the first from "Old Greece" to enter the Roman Senate, as representatives of the two "great powers" of the Classical Age. The Temple of Olympian Zeus had been under construction for more than five centuries; Hadrian committed the vast resources at his command to ensure that the job would be finished.

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He also organised the planning and construction of a particularly challenging and ambitious aqueduct to bring water to the Athenian agora.

Coins celebrate him as the restorer of the island. In early March Hadrian set off on a tour of Italy; his route has been reconstructed through the evidence of his gifts and donations. Less welcome than such largesse was his decision in to divide Italy into four regions under imperial legates with consular rank, acting as governors.

They were given jurisdiction over all of Italy, excluding Rome itself, therefore shifting Italian cases from the courts of Rome. His arrival coincided with the good omen of rain, which ended a drought. Along with his usual role as benefactor and restorer, he found time to inspect the troops; his speech to them survives.