Uncles ex Machina: Familial Epiphany in Euripides' Electra

At the close of Euripides’ Electra, the Dioscuri suddenly appear ‘on high’ to their distraught niece and nephew, who have just killed their mother, the divine twins’ mortal sister. This is in fact the second longest extant deus ex machina (after the final scene in Hippolytus), and the only scene in which a tragedian attempts to resolve directly the aftermath of the matricide. In this article, I argue that Castor's and Polydeuces’ sudden apparition to Orestes and Electra constitutes a specialised point of intersection between the mortal and immortal realms in Greek tragedy: familial epiphany, an appearance by a god who has an especially intimate relationship with those on stage. Euripides’ focus on the familial divine as a category accentuates various contradictions inherent to both ancient Greek theology and dramaturgy. The Dioscuri are a living paradox, ambiguously traversing the space between dead heroes and gods, managing at the same time to occupy both. They oscillate uniquely between the mortal and immortal worlds, as different sources assign different fathers to each brother, and others speak of each one possessing divinity on alternate days. As I propose, the epiphany of these ambiguous brothers crystallises the problem of the gods’ physical presence in drama. Tragedy is the arena in which gods burst suddenly into the mortal realm, decisively and irrevocably altering human action. The physical divine thus tends to be both marginal and directorial, tasked with reining in the plot or directing its future course. The appearance of the familial divine, on the other hand, can in fact obscure the resolution and future direction of a play, undermining the authority of the tragic gods. In the specific case of Electra, I contend that the involvement of the Dioscuri, who are Electra's and Orestes’ maternal uncles, produces a sense of claustrophobia at the close of the play, which simultaneously denies the resolution that is expected from a deus ex machina while also revealing the pessimistic nature of what is typically considered a reassuringly ‘domestic’ and character driven drama

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-0

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>antibodies against IκBα and p65/RelA. A secondary antibody conjugated with Texas Red (Molecular Probes) was used. Images were taken by confocal microscopy

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-4

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>expression vectors or (b) pcDNA3.1 and/or CMV-Tat and/or CMV-IκBα expression vectors, as indicated. Cells were treated with LMB immediately after transfection and/or with PMA two hours after transfection, as indicated. Luciferase activity was measured 18 hours after transfection. Numbers on the top of the bars represent fold transcriptional activity relative to unstimulated T cells transfected with pcDNA3.1

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-8

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>t with LMB up to 30 minutes. Photographs were taken by confocal microscopy every minute after adding LMB

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-3

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>�B/IκBα complexes in the nucleus after treatment with LMB or PMA. Ten micrograms of cytosolic and nuclear extracts from CDT cells treated with either PMA or LMB during 4 and 6 hours respectively were analyzed by Western Blot using antibodies against p65/RelA and IκBα. Immunoprecipitation assays were performed using 100 μg of these cytosolic and nuclear extracts, which were incubated with 5 μg of an antibody against p65/RelA conjugated with agarose. IκBα and p65/RelA complexes were characterized by immunoblotting. (b) Contamination with cytosolic proteins during nuclear protein extraction or accumulation of cytosolic proteins in the nucleus after treatment with LMB was assessed by Western Blot using an antibody against both p105 and p50/NF-κB1 proteins. (c) Analysis of NF-κB DNA-binding activity in CDT cells treated with either PMA or LMB. Three micrograms of nuclear extract were incubated with an oligonucleotide containing the double consensus motif κB present in the HIV LTR labeled with [α-P]-dCTP. Protein extracts were obtained from CDT cells after treatment with either LMB or PMA for 6 and 4 hours respectively. (d) Analysis of IκBα pool dependence on protein synthesis. Ten micrograms of nuclear extracts from CDT cells incubated with 20 nM LMB for 4 hours and 10 μg/ml CHX and/or 25 ng/ml PMA for 4 hours,30 min and 2 hours, respectively, were analyzed by Western Blot

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-2

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>t with LMB up to 30 minutes. Photographs were taken by confocal microscopy every minute after adding LMB

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-5

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>together with CMV-IκBα or pcDNA3.1 as negative control, and then activated with anti-CDand IL-2, PHA and IL-2, or maintained in the absence of activation. Viral replication was determined by quantification of HIV p24-gag antigen in culture supernatants (a) after 5 days of transfection or (b) after 7 days of transfection. Numbers on the top of the bars represent fold HIV-replication relative to unstimulated T cells transfected with pcDNA3.1. Differences in p24-gag production were significant for resting and anti-CD-activated T cells (p < 0.05) and a trend towards statistical significance was found in PHA-activated T cells (p = 0.081)

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-1

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>ctors per million of cells. LMB was added immediately after transfection. After 18–24 hours of incubation, cells were analyzed by confocal microscopy. pEYFP-C1 vector was used as control of unspecific distribution. (b) Resting purified CDT lymphocytes were transiently transfected with the control plasmid pcDNA3.1 by using an Amaxa nucleofector and a classical electroporator (Equibio). As occurs in untransfected resting T cells (lane 1), NF-κB was not induced in resting CDT lymphocytes after electroporation (lanes 3 and 4). As a positive control, NF-κB (p50/p65) binding was induced in these cells by PMA activation (lane 2)

Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression-7

<p><b>Copyright information:</b></p><p>Taken from "Basal shuttle of NF-κB/IκBα in resting T lymphocytes regulates HIV-1 LTR dependent expression"</p><p>http://www.retrovirology.com/content/4/1/56</p><p>Retrovirology 2007;4():56-56.</p><p>Published online 8 Aug 2007</p><p>PMCID:PMC1988826.</p><p></p>ctors per million of cells. LMB was added immediately after transfection. After 18–24 hours of incubation, cells were analyzed by confocal microscopy. pEYFP-C1 vector was used as control of unspecific distribution. (b) Resting purified CDT lymphocytes were transiently transfected with the control plasmid pcDNA3.1 by using an Amaxa nucleofector and a classical electroporator (Equibio). As occurs in untransfected resting T cells (lane 1), NF-κB was not induced in resting CDT lymphocytes after electroporation (lanes 3 and 4). As a positive control, NF-κB (p50/p65) binding was induced in these cells by PMA activation (lane 2)

Analysis of caspase-3 activation in TCERG1-depleted cells after staurosporine induction.

<p>(A) Activation of caspase-3 was decreased in TCERG1-knockdown Jurkat cells treated with staurosporine. Caspase-3 activation was measured by chemiluminescence in control and TCERG1-knockdown Jurkat cells treated (+STAU) or not with 0.5 μM staurosporine (-STAU) for 1 h. The bar diagram shows the fold activation from four independent experiments (means ± SEM). Data obtained from cells without staurosporine were set as 1. *, <i>p</i> < 0.05. (B) Analysis of TCERG1, procaspase-3 cleavage (precursor p35 and cleaved fragments p20/p17), and PARP-1 cleavage (p89) levels. Protein expression was analyzed by immunoblotting using specific antibodies in protein extracts obtained from control and TCERG1-knockdown Jurkat cells treated (+STAU) or not with 0.5 μM staurosporine (-STAU) for 1 h. ß-actin was used as a loading control. The quantification of the bands is shown below each panel.</p