Romanian Biotechnological Letters Vol. 18, No.4, 2013
Copyright © 2013 University of Bucharest Printed in Romania. All rights reserved
ORIGINAL PAPER
Romanian Biotechnological Letters, Vol. 18, No. 4, 2013 8539
Current methods for investigation of the unfolded protein response
Received for publication, March 15, 2013
Accepted, July 15, 2013
ALINA STROESCU, LIVIA SIMA, FLORENTINA PENA*
Institute of Biochemistry of the Romanian Academy, Splaiul Independentei 296, 060031
Bucharest 17, Romania.
* Corresponding author (Email: Florentina.Pena@biochim.ro; phone: +40212239069; fax
+40212239068)
Abstract
The endoplasmic reticulum (ER) is the cellular organelle where proteins from the secretory
pathway fold. The ER is a very dinamic organelle that can respond rapidly to changes in the ER load by
upregulating the ER resident proteins that are involved in the folding process. Simultaneously, less
proteins from the secretory pathway are synthesized. The whole cascade of events is called the unfolded
protein response (UPR). The UPR can originate from physiological or pathological conditions and
many times the ER changes in a subtle manner. Researchers working in the ER field developed various
tools to assay the ER stress. Here we describe methods optimized in our laboratory and show that
western blotting, quantitative real time polymerase chain reaction (RT-qPCR) and reverse
transcription-polymerase chain reaction (RT-PCR) are reliable, cost-effective techniques that
researchers can choose to evidence ER stress activation.
Abbreviations:
BiP, immunoglobulin heavy-chain binding protein; ER, endoplasmic reticulum; TG, thapsigargin; TN,
tunicamycin; UPR, unfolded protein response; IRE1, inositol-requiring kinase 1; XBP1, X-box-
binding protein 1; PERK, RNA-dependent protein kinase-like ER kinase; ATF6, activating
transcription factor 6; eIF2 , eukaryotic translational initiation factor 2 Bcl-2, B cell lymphoma
2, ATF4, activating transcription factor 4; CHOP, CCAAT/enhancer-binding protein-homologous
protein; GRP, glucose-regulated proteins, CREB-RP, cyclic AMP-response-element-binding protein-
related protein; S1P, site-1 protease;S2P, site-2 protease; ERSE, ER stress response elements; Grp94,
Glucose-regulated protein 94; ERp44, endoplasmic reticulum resident protein 44 kDa; ERp72,
endoplasmic reticulum resident protein 72 kDa; ERp57, endoplasmic reticulum resident protein 57
kDa; Ero1 , ER oxidoreductin 1 ; DTT , dithiothreitol; GAPDH, Glyceraldehyde-3-Phosphate
Dehydrogenase; ERAD, ER-associated degradation; ECL, enhanced chemiluminescence; RT-PCR,
reverse transcription-polymerase chain reaction; RT-qPCR, quantitative real time polymerase chain
reaction.
1. The endoplasmic reticulum – a short overview
The endoplasmic reticulum (ER) is essential for production of secretory proteins
(BRAAKMAN [1]). Proteins from secretory pathway emerge from ER-bound ribosomes and
are further modified and folded by the ER folding machinery. There are three major classes of
ER resident proteins: molecular chaperones, foldases and lectins (SCHRODER [2]).
The molecular chaperones facilitate protein folding by shielding unfolded regions
from surrounding proteins. BiP for instance binds to the hydrophobic patches that are exposed
during folding, thus preventing aggregation. The foldases (cis–trans peptidyl–prolyl
isomerases, PPI/immunophilins and PDIs) catalyze steps in protein folding such as the
formation and isomerization of disulfide bonds. Lectins interact with glycoproteins due to the
presence of a lectin site that can recognize an early folding oligosaccharide processing
intermediate on the folding glycoprotein (WILLIAMS [3]). Correctly folded secretory
proteins are exported to intracellular organelles or to the extracellular surface. Proteins