J-Perk 5.01 serial key or number
J-Perk 5.01 serial key or number
PMC
B. L. Williams
Jerome L. and Dawn Greene Infectious Disease Laboratory, Mailman School of Public Health, Columbia University, New York, New York
W. I. Lipkin
Jerome L. and Dawn Greene Infectious Disease Laboratory, Mailman School of Public Health, Columbia University, New York, New York
Abstract
Borna disease virus infection of neonatal rats results in a characteristic behavioral syndrome and apoptosis of subsets of neurons in the hippocampus and cerebellum (neonatal Borna disease [NBD]). The cellular mechanisms leading to neurodevelopmental damage in NBD have not been fully elucidated. Insights into this model may have general implications for understanding the pathogenesis of virus-associated neurodevelopmental damage. Here we report the presence of endoplasmic reticulum (ER) stress markers and activation of the unfolded protein response in the NBD hippocampus and cerebellum. Specific findings included enhanced PERK-mediated phosphorylation of eif2α and concomitant regulation of ATF4 translation; IRE1-mediated splicing of XBP1 mRNA; and cleavage of the ATF6 protein in NBD rat brains. We found evidence for regional and cell type-specific divergence in the expression of ER stress-induced proapoptotic and quality control signals. Our results demonstrate that ER stress induction in death-susceptible Purkinje neurons in NBD is associated with the expression of the proapoptotic molecule CHOP in the absence of compensatory expression of the ER quality control molecules Bip and protein disulfide isomerase. In contrast, ER stress in death-resistant astrocytes is associated with complementary expression of CHOP and ER quality control signals. These results implicate an imbalance between ER stress-mediated apoptosis and survival signaling as a critical determinant of neural cell fate in NBD.
Borna disease virus (BDV) is a nonsegmented, negative-sense, single-stranded RNA virus belonging to the virus family Bornaviridae within the order Mononegavirales (5, 11). Experimental infection of adult immunocompetent Lewis rats results in a marked CD8+ immune response (meningoencephalitis) and a progressive movement disorder (20). In contrast, newborn rats infected with BDV (neonatal Borna disease [NBD]) do not mount an overt cellular immune response yet have prominent astrogliosis and microgliosis, altered cytokine, neurotrophic factor, and neurotrophic factor receptor gene expression, abnormal development of brain monoaminergic systems, cerebellar and hippocampal dysgenesis, and disturbances of learning, mood, and behavior (8, 25, 47, 54). Although BDV is noncytolytic, NBD is attended by apoptotic degeneration of neurons in the cerebellum (CBLM) and hippocampus (HC) that undergo substantial postnatal maturation (25, 72). However, the cellular and molecular mechanisms leading to neurodegeneration in NBD remain unclear.
The endoplasmic reticulum (ER) plays a vital role in a variety of cellular functions, including synthesis, folding, and posttranslational modification of secretory and membrane proteins, cytoplasmic and mitochondrial metabolism, homeostasis of intracellular calcium, and apoptosis. The ER is sensitive to nutrient or energy deprivation, disruption of calcium homeostasis, inhibition of N-linked glycosylation, oxidative stress, and accumulation of unfolded, misfolded, or excessive protein (61, 69). Perturbations causing ER stress result in accumulation of unfolded proteins in the ER lumen and initiation of a complex signal transduction cascade, known as the unfolded protein response (UPR) (Fig. (Fig.1).1). The UPR is activated to restore the ER to its normal physiological state by two adaptive mechanisms: (i) increasing the folding capacity of the ER through induction of molecular chaperones and foldases and (ii) decreasing protein burden on the ER through global inhibition of translation and enhancing degradation and clearance of unfolded proteins. Essential to this quality control mechanism is the ATP-dependent ER chaperone, Bip (immunoglobulin heavy chain-binding protein). In the physiological state, Bip binds to and suppresses the activity of three proximal, ER-resident, transmembrane, UPR transducers: PKR-like ER kinase (PERK), inositol-requiring enzyme 1 (IRE1), and activating transcription factor 6 (ATF6).
Schematic illustration of ER stress and UPR activation. ER stresses can result in accumulation of unfolded proteins in the ER. Unfolded proteins are bound by Bip, resulting in release and activation of three ER-resident transmembrane transducers of the UPR: PERK, IRE1, and ATF6. Activated PERK phosphorylates eif2α, attenuating global protein synthesis, and enhances translation of ATF4. Activated IRE1 facilitates splicing of XBP1U, resulting in translation of the transcriptionally active form of XBP1 (XBP1S). Following Bip release, ATF6 translocates to the Golgi, where it is cleaved by proteases, releasing the transcriptionally active form of ATF6 (ATF6p50). ATF4, XBP1S, and ATF6p50 enter the nucleus, where they activate the expression of a distinct set of UPR target genes.
During ER stress, Bip binds misfolded proteins and is thereby sequestered from PERK, IRE1, and ATF6 (3). When released from Bip, the kinase domain of activated PERK phosphorylates Ser51 of the alpha subunit of eukaryotic translation initiation factor 2 (eif2α), thereby attenuating global translation initiation and protein synthesis (17). Paradoxically, PERK-mediated phosphorylation of eif2α preferentially induces translation of activating transcription factor 4 (ATF4) (18). Following Bip release, IRE1 undergoes dimerization and trans-autophosphorylation, activating its endoribonuclease activity; IRE1 then mediates the excision of a 26-nucleotide intron from X-box binding protein 1 (XBP1), resulting in a translational frameshift and formation of a potent transcriptional activator (78). Bip-free ATF6 is transported to the Golgi, where it is cleaved by site-1 and site-2 proteases, generating a 50-kDa bZIP transcription factor (ATF6p50) that migrates to the nucleus (22, 62, 75). The downstream nuclear effectors of the UPR (ATF4, spliced XBP1, and ATF6p50) regulate the expression of genes that enhance the protein folding and degrading capacity of the ER, regulate amino acid metabolism, mediate resistance to oxidative stress, and induce apoptosis (19, 35, 38).
The ER elicits apoptotic signals when protein load exceeds its folding capacity. C/EBP homologous protein (CHOP), also known as GADD153 (growth arrest- and DNA damage-inducible gene 153), is induced by ER stress. Overexpression of CHOP triggers cell cycle arrest and apoptosis, down-regulates the prosurvival molecule Bcl-2, and promotes the production of reactive oxygen species (1, 44-46, 49). In contrast, overexpression of the ER chaperone Bip reduces CHOP induction associated with ER stress and attenuates apoptosis (71). Thus, the cellular fate with ER stress reflects the coordinated interaction of quality control functions and proapoptotic signals. ER stress induction and regulation have been reported following in vitro infections by several viruses, including hepatitis C virus (HCV), Japanese encephalitis virus, bovine viral diarrhea virus, respiratory syncytial virus, human cytomegalovirus (HCMV), and Tula hantavirus (4, 26, 27, 39, 63, 65).
Here we demonstrate that BDV infection of newborn rats results in neural and astrocytic ER stress, induces the UPR, and results in a neural cell-specific imbalance between proapoptotic and survival signaling.
MATERIALS AND METHODS
Animals and virus inoculation.
Lewis rat dams were obtained from Charles River Laboratories (Wilmington, MA). Within 12 h of birth, 50 Lewis rat pups were inoculated into the right cerebral hemisphere with 50 μl of 5 × 103 tissue culture infectious units of BDV strain He/80-1 (NBD) or phosphate-buffered saline (control: sham inoculated). Rats were sacrificed at postnatal days 21, 28, and 84 (PND21, PND28, and PND84) for nucleic acid, protein, and anatomic analyses.
RNA extraction.
At PND28 postinoculation, NBD (n = 7) and control (n = 5) rats were terminally anesthetized with CO2. CBLM and HC were immediately dissected, snap frozen in TRIzol (Invitrogen, Carlsbad, CA), and stored at −80°C. Following extraction using standard protocols, RNA was quantitated by UV spectrophotometry.
DNA microarray analysis.
Equal amounts of total RNA extracted from either CBLM or HC of individual PND28 animals were pooled to create a single control RNA sample (n = 5 control rats) and an NBD RNA pool (n = 7 NBD rats) for use in DNA microarray assays. Indirect labeling and cDNA synthesis were performed using 5 μg of pooled RNA and the 3DNA Array 350 expression array detection kit for microarrays (Genisphere, Hatfield, PA). Cy3 and Cy5 capture sequence-labeled products were hybridized to a rat 8,000-gene oligonucleotide array (Public Health Research Institute, Newark, NJ) in sodium dodecyl sulfate (SDS)-based hybridization buffer containing denatured human Cot-1 (1 μg/40 μl; Invitrogen/Gibco LTI, Gaithersburg, MD) for 18 h at 53°C. Posthybridization washes were followed by an additional hybridization with fluorescent Cy3-3DNA and Cy5-3DNA capture reagents in SDS hybridization buffer for 3 h at 62°C, followed by sequential washes. Fluorescence-labeled slides were dried by centrifugation and scanned with a GenePix 4000B scanner (Axon Instruments, Inc., Union City, CA) at 5-μm resolution. For each spot, background-subtracted signal intensity, acquired using GenePix Pro 4.1 software, was normalized for both Cy3 and Cy5 expression by scaling individual spot intensities in each channel to the total intensity of all spots on the array. Experiments using CBLM or HC RNA sets were performed in triplicate. Data were paired for each array and analyzed using the Cyber T web interface (http://visitor.ics.uci.edu/genex/cybert/). Differences in gene hybridization signal that met the following criteria were considered to be significant: n-fold change greater than 1.5 or less than −1.5 and P value of <0.05 (paired t test).
Quantitative SYBR Green real-time PCR.
Intron/exon spanning, gene-specific PCR primers specific for rat CHOP and porphobilinogen deaminase (PBGD) as a housekeeping gene control were designed for real-time PCR using Primer Express 1.0 software (Applied Biosystems, Foster City, CA). Published primers were used for XBP1 and Bip assays (Table (Table1)1) (36, 51). PCR standards for determining copy numbers of target transcripts were cloned into the vector pGEM-T easy (Promega Corporation, Madison, WI). Linearized plasmids were quantitated by UV spectroscopy, and 10-fold serial dilutions were created in water containing yeast tRNA (1 ng/μl). RNA from CBLM or HC of individual animals was used for real-time PCR assays. cDNA was synthesized using Taqman reverse transcription reagents (Applied Biosystems) from 2 μg unpooled RNA per 100 μl reaction mixture from either the CBLM or HC of each of seven NBD rats and five control rats; each sample was assayed in triplicate. Each 25-μl amplification reaction mixture contained 10 μl template cDNA, 12.5 μl SYBR Green Master Mix (Applied Biosystems), and 300 nm gene-specific primers (Table (Table1).1). The thermal cycling profile using a Model 7700 sequence detector system (Applied Biosystems) consisted of the following: stage 1, 1 cycle at 50°C for 2 min; stage 2, 1 cycle at 95°C for 10 min; stage 3, 45 cycles at 95°C for 15 s and 60°C for 1 min (for CHOP amplification); stage 1, 1 cycle at 50°C for 2 min; stage 2, 1 cycle at 95°C for 10 min; stage 3, 45 cycles at 94°C for 30 s, 55°C for 30 s, and 72°C for 30 s (for Bip amplification); and stage 1, 1 cycle at 50°C for 2 min; stage 2, 1 cycle at 95°C for 10 min; stage 3, 45 cycles at 94°C for 1 min, 60°C for 1 min, and 72°C for 1 min (for XBP1 amplification). A PBGD fragment was amplified in triplicate reactions by real-time PCR on the same plate as the gene of interest. The mean concentration of PBGD in each sample was used to control for integrity of input RNA and to normalize values of target gene expression to those of the housekeeping gene expression. The final results were expressed as the mean number of copies per 200 ng total RNA for CHOP, Bip, or XBP1, relative to values obtained for PBGD RNA.
Western blot analysis.
Individual CBLM and HC were dissected from PND28 NBD (n = 4) and control (n = 4) rats and homogenized in ice-cold hypotonic cell lysis buffer (10 mM HEPES, pH 7.9, 10 mM KCl, 0.1 mM EDTA, 0.1 mM EGTA) containing protease inhibitors (Complete Mini, EDTA-free tablets; Roche Molecular Biochemicals, Indianapolis, IN) and the serine-threonine phosphatase inhibitor β-glycerophosphate (17.5 mM, EMD Biosciences, San Diego, CA) and incubated on ice for 15 min. Homogenates were centrifuged at 8,000 × g for 15 min at 4°C. Supernatants containing cytoplasmic proteins were collected, and protein concentrations estimated by Bradford assay (Bio-Rad, Hercules, CA). Cell pellets were washed five times with cell lysis buffer and incubated for 30 min on ice with nuclear lysis buffer (20 mM HEPES, pH 7.9, 400 mM NaCl, 1 mM EDTA, 1 mM EGTA, and 1 mM dithiothreitol) containing protease and phosphatase inhibitors, followed by centrifugation for 20 min at 15,000 × g. Cleared supernatants containing nuclear extracts were collected and protein concentration estimated by Bradford assay (Bio-Rad). Cytoplasmic or nuclear proteins from regional brain homogenate (20 μg) in sample buffer (10 mM Tris-Hcl, pH 7.5, 10 mM EDTA, 20% [vol/vol] glycerol, 1% [wt/vol] SDS, 0.005% [wt/vol] bromphenol blue, 100 mM dithiothreitol) were boiled for 5 min and size fractionated by 10% SDS-polyacrylamide gel electrophoresis. SYPRO Ruby staining of cytoplasmic proteins was carried out according to the manufacturer's protocol (Invitrogen, Carlsbad, CA). Proteins were transferred to nitrocellulose membranes using a semidry blotting apparatus (Owl Separation Systems, Portsmouth, NH). Membranes were blocked in 2% nonfat milk powder (for CHOP, ATF4, ATF6, XBP1, and glyceraldehyde phosphate-3-dehydrogenase [GAPDH]) or 2% bovine serum albumin (for phospho-eif2α) in TTBS (20 mM Tris-Hcl, pH 7.6, 137 mM NaCl, 0.1% Tween 20) overnight at room temperature and incubated with mouse monoclonal anti-CHOP (1:100; Santa Cruz Biotechnology), mouse monoclonal anti-PDI (1:500; Affinity BioReagents, Golden, CO), rabbit anti-XBP1 (1:100; M-186; Santa Cruz Biotechnology), mouse monoclonal anti-ATF6 (1:200; IMGENEX, San Diego, CA), or rabbit anti-ATF4 (1:50; C-20; Santa Cruz Biotechnology) in TTBS with 1% nonfat milk or rabbit monoclonal anti-phospho-eif2α (Ser51) (1:300; Cell Signaling Technology, Danvers, MA) in TTBS with 2% bovine serum albumin for 2 h at room temperature. Membranes were washed three times for 10 min each with TTBS prior to incubation with peroxidase-conjugated goat antimouse immunoglobulin G (IgG) (1:2,000; Bio-Rad) or goat antirabbit IgG (1:2,000; Bio-Rad) in TTBS with 1% nonfat dry milk for 1 h at room temperature. Membranes were developed using the ECL Western blot detection system (Amersham Biosciences, Arlington Heights, IL) and scanned for chemiluminescence using a Storm 840 imager (Molecular Dynamics, Sunnyvale, CA). Blots were stripped and reprobed with mouse anti-GAPDH monoclonal antibody (Ambion, Austin, TX), as a housekeeping gene, for normalization of cytoplasmic proteins. Protein bands were quantified using Image Quant software (v.1.0; Molecular Dynamics).
XBP1 RT-PCR splicing analysis.
ER stress-induced processing of XBP1 mRNA was evaluated by PCR and restriction site analysis. Total RNA (2 μg) extracts from CBLM and HC of individual PND28 NBD (n = 7) and control (n = 5) rats were reverse transcribed using MultiScribe reverse transcriptase (RT) and random hexamer primers (Applied Biosystems, Foster City, CA). The 601-bp PCR product, encompassing the IRE1 cleavage site of XBP1, was amplified using the following XBP1 primers: upper strand primer, 5′-AAACAGAGTAGCAGCGCAGACTGC-3′; and lower strand primer, 5′-GGATCTCTAAAACTAGAGGCTTGGTG-3′. The thermal cycling profile consisted of 30 cycles at 94°C for 1 min, 60°C for 1 min, and 72°C for 1 min. PCR products were either directly resolved on a 2% agarose gel or incubated with PstI at 37°C for 5 h followed by separation on 2% agarose gels. Bands representing spliced (XBP1S) and unspliced (XBP1U) forms of XBP1 from seven NBD rats were extracted from agarose and subcloned into pGEM-T Easy Vector (Promega Corporation) for sequencing.
Histological analysis and immunofluorescence.
Under CO2 anesthesia, PND21, PND28, and PND84 NBD (n = 5/age group) and control (n = 4/age group) rats were perfused via left ventricular puncture with phosphate-buffered saline (1 ml/g body weight), followed by buffered 4% paraformaldehyde (1 ml/g body weight). Brains were postfixed in 4% paraformaldehyde overnight at 4°C and cryoprotected with graded sucrose solutions. Cryostat sections (14 μm) were collected onto glass slides (Super Frost Plus; Fisher Scientific, Pittsburgh, PA). Tissue sections were stained with hematoxylin and eosin (H&E) for histological analysis. Immunofluorescence microscopy was carried out as previously described (74), using the following primary antibodies: rabbit anti-BDVp40 (nucleoprotein; 1:1500) (6), mouse monoclonal anti-CHOP (1:50; Santa Cruz), rabbit anti-P-eif2α (Ser51) monoclonal antibody (1:50; Cell Signaling), rabbit anti-ATF4 (1:50; Santa Cruz), rabbit anti-Bip (1:50; Santa Cruz), mouse monoclonal anti-PDI (1:500; Affinity BioReagents), rabbit anti-glial fibrillary acidic protein (GFAP) antibody (1:100; DAKO Cytomation, Carpinteria, CA), mouse anti-GFAP monoclonal antibody cocktail (1:30; BD Pharmingen, San Diego, CA), rabbit anti-calbindin D-28K (1:100; Chemicon, Temecula, CA). Secondary antibodies were Cy3-conjugated antimouse or antirabbit IgG (1:200; Jackson Immunoresearch Laboratories, Inc., West Grove, PA) and/or Cy2-conjugated antimouse or antirabbit IgG (1:200; Jackson Immunoresearch).
Statistical analysis.
The significance of observed differences among NBD and control groups was assessed by paired t test for microarray experiments or analysis of variance (ANOVA) for real-time PCR and Western immunoblot analysis. Analysis was carried out using StatView software (v.5.0.1; SAS Institute Inc., Cary, NC). Values were considered to be significant when P values were <0.05.
RESULTS
Microarray analysis reveals differential expression of ER stress genes in NBD.
Intracranial inoculation of Lewis rats with BDV within the first 24 h of life results in persistent infection, behavioral deficits, and progressive apoptosis of neurons that undergo postnatal maturation. To identify specific molecular and cellular pathways that mediate these effects, we profiled mRNA expression in brains of PND28 rats, a time point that corresponds with peak histologic evidence of apoptotic neural damage in CBLM and HC (25, 72).
Microarray analysis of mRNA extracted from the CBLM and HC of PND28 NBD and control rats revealed altered mRNA levels for 11 genes previously reported to be induced by ER stress and one ER stress-repressed gene (Fig. (Fig.2A)2A) (hereafter referenced as group 1 genes) (19, 35, 48, 56, 60). In some instances, genes met microarray criteria in either CBLM or HC but not both; where degrees of difference were modest, a trend toward altered gene expression was evident in both regions. Of the group 1 genes, five met microarray criteria in both NBD CBLM and HC (CHOP, PDI, Herp [homocysteine-induced ER protein], NEFA [novel DNA-binding/EF-hand/leucine zipper protein], and Slc3a2 [4F2 antigen heavy chain-amino acid transporter]); six specifically met criteria in CBLM (Asn-S [asparagine synthetase], Hmox1 [heme oxygenase], gas-5 [growth arrest-specific 5], GP [glycogen phosphorylase], Gadd45 [growth arrest and DNA-damage-inducible 45], and HMG-CoA-S [3-hydroxy-3-methylglutaryl-coenzyme A synthase]); and only one gene specifically met criteria in the HC (DRAL [down-regulated in rhabdomyasarcoma LIM]). Microarray experiments also identified alterations in genes involved in UPR activation (PERK) or reglucosylation of nonnative glycoproteins in the ER (UGGT [UDP-glucose-glycoprotein glucosyltransferase]) (Fig. (Fig.2A)2A) (hereafter referenced as group 2 genes) and ER-resident calcium transporters (RyR1 [ryanodine receptor type 1], RyR3 [ryanodine receptor type 3], SERCA2 [sarcoplasmic reticulum Ca2+-ATPase 2], and SERCA3 [sarcoplasmic reticulum Ca2+-ATPase 3]) (Fig. (Fig.2A;2A; hereafter referenced as group 3 genes) (58, 69). Interestingly, all four calcium transporters specifically met microarray criteria in the NBD CBLM, with increased expression of RyR1 and RyR3 and decreased expression of SERCA2 and SERCA3. n-fold change and P values for these genes are given for comparison (Fig. (Fig.2B2B).
Identification of ER stress genes in PND28 NBD rats by microarray analysis. Microarray analysis was conducted using pooled total RNA isolated from dissected CBLM and HC of PND28 NBD (n = 7) and PND28 control (n = 5) rats. (A) Results represent mean n-fold change ± standard deviation for genes from three independent experiments comparing RNA from NBD CBLM versus control CBLM (black bars) and mean n-fold change ± standard deviation for genes in NBD HC (gray bars) versus control HC. Genes were considered significant when n-fold induction or repression in NBD rats was >1.5 or <−1.5 and the P value was <0.05 compared to control results. Black asterisks denote genes meeting these criteria in NBD CBLM; gray asterisks denote genes meeting these criteria in NBD HC. Genes are grouped into known ER stress-responsive genes (group 1), genes involved in ER stress response (group 2), and regulators of ER calcium homeostasis (group 3). (B) Table showing n-fold induction or fold repression and P values for ER stress genes (accession no. provided) for NBD rat CBLM and HC. For gene abbreviations, see the text.
Phosphorylation of eif2α in NBD rat brain and increased translation of ATF4.
ER stress in mammalian cells leads to the activation of PERK. Activated PERK inhibits general protein biosynthesis through phosphorylation of eif2α, limiting further protein overload in the ER (17). Activation of the PERK pathway in PND28 NBD rats was assessed in Western blots of cytoplasmic extracts of CBLM and HC using antibodies that bind the phosphorylated form of eif2α (P-eif2α) and GAPDH (loading control). P-eif2α antibodies detected a single band of 40 kDa corresponding to the expected size of P-eif2α in both control and NBD CBLM and HC extracts (Fig. (Fig.3A).3A). Quantitation of P-eif2α band densities normalized to GAPDH revealed a 1.36-fold increase in P-eif2α in NBD CBLM (ANOVA, P = 0.0136) (Fig. (Fig.3B).3B). Levels of P-eif2α were increased 2.67-fold in NBD HC (ANOVA, P = 0.0007) (Fig. (Fig.3C3C).
Activation of the PERK/eif2α/ATF4 arm of the UPR in NBD rats. (A) Cytoplasmic lysates (20 μg) from CBLM and HC of control and NBD rats were evaluated by Western analysis using antibodies specific for the phosphorylated form of eif2α (P-eif2α; upper panels) relative to levels of GAPDH (housekeeping gene; lower panels). (B and C) Determination of P-eif2α band density relative to that for GAPDH from control (n = 4) or NBD (n = 4) rat CBLM (B) (1.36-fold increase in NBD; ANOVA, P = 0.0136) or HC (C) (2.67-fold increase in NBD; ANOVA, P = 0.0007). (D) Sypro Ruby stain of protein extract (20 μg) from CBLM and HC of control and NBD rats. Note that relative levels of proteins are similar between control and NBD rats in both brain regions, with the exception of a single band in NBD lysates from HC corresponding to the BDV nucleoprotein (arrowhead). (E) Cytoplasmic (upper panel) and nuclear (lower panel) lysates from CBLM of control and NBD rats were evaluated by Western analysis using antibodies specific for ATF4. Note equal amounts of ATF4 protein in cytoplasmic lysates and higher ATF4 band signal in nuclear extracts from NBD rats compared to results for controls. (F) Determination of ATF4 band density from control (n = 4) or NBD (n = 4) nuclear protein extracts revealed increased ATF4 levels in NBD CBLM (3.06-fold increase in NBD; ANOVA, P = 0.0223). Asterisks indicate P values of <0.05.
Phosphorylation of eif2α prevents the formation of the ternary translation initiation complex (eif2/GTP/Met-tRNAiMet), thereby attenuating global protein synthesis (17). We assessed total protein levels in CBLM and HC from control and NBD rats using SYPRO Ruby staining of SDS-polyacrylamide gel electrophoresis gels loaded with equal amounts of cytoplasmic protein extracts. No differences were detected between NBD and controls in either CBLM or HC except for the presence of a protein band in NBD subsequently identified by Western blot analysis as the BDV nucleoprotein (Fig. (Fig.3D,3D, arrowhead).
PERK-mediated phosphorylation of eif2α inhibits general protein translation but selectively promotes translation of ATF4 mRNA during ER stress. In unstressed cells, upstream open reading frames in the 5′ untranslated region of ATF4 mRNA repress the translation of ATF4. During ER stress, eif2α phosphorylation allows ribosomal bypass scanning of the ATF4 mRNA upstream open reading frames, resulting in efficient ATF4 translation (18). ATF4 activates the transcription of genes involved in amino acid metabolism and transport (i.e., genes encoding Asn-S and Slc3a2), oxidative stress responses (i.e., that encoding Hmox-1), and ER stress-induced apoptosis (i.e., that encoding CHOP) (19). Given microarray results indicating increased levels of transcripts corresponding to ATF4-regulated genes in NBD, we assessed levels of the ATF4 protein in cytoplasmic and nuclear extracts of PND28 NBD and control CBLM and HC. ATF4 (26 kDa) levels were 3.06-fold higher in nuclear extracts of NBD CBLM (ANOVA, P = 0.0223) (Fig. 3E and F). No significant differences in ATF4 were observed between NBD and the controls in cytoplasmic or nuclear extracts from HC (data not shown).
XBP1 splicing in NBD rats.
ER stress results in Bip release from IRE1 and activation of IRE1 RNase activity. XBP1 mRNA serves as a specific substrate for IRE1 RNase (78). Thus, splicing of XBP1 is commonly used as a marker of IRE1 activation. We assessed the activation state of IRE1 by investigating XBP1 mRNA splicing and expression, as well as XBP1 protein expression, in PND28 NBD and control rats.
XBP1 splicing was evaluated by RT-PCR using primers that amplify products spanning the 26-nucleotide excision site. Agarose gel analysis of XBP1 cDNA amplified from CBLM (Fig. (Fig.4A)4A) and HC (Fig. (Fig.4B)4B) mRNA of individual controls (n = 5) revealed a single, abundant product of approximately 601 bp, the expected amplicon size of unspliced XBP1 (XBP1U). In NBD, an additional lower-molecular-weight band of approximately 575 bp (the expected size of XBP1S) was observed in CBLM and HC (Fig. 4A and B, respectively). Splicing of XBP1 disrupts a PstI restriction site localized within the 26-nucleotide excision site. Thus, PstI digestion is expected to completely cut XBP1U into two fragments of 312 bp and 289 bp, while XBP1S should be resistant to digestion. As expected, XBP1 PCR products were completely digested by PstI in CBLM and HC of all controls (Fig. 4C and D, respectively) while XBP1S was resistant to PstI digestion in all NBD rats (Fig. 4C and D). Sequencing of gel-extracted PCR fragments from NBD and control rats confirmed that the PCR products represented the expected sequences for the XBP1U and XBP1S forms of XBP1 (Fig. (Fig.4E4E).
XBP1 mRNA is selectively spliced in NBD rats. RNA extracted from CBLM and HC of PND28 control (n = 5) and NBD (n = 7) rats was analyzed by RT-PCR using primers that amplify both unspliced XBP1 (XBP1U) and spliced (XBP1S) XBP1. (A and B) Representative gel images of XBP1 RT-PCR. Note detection of XBP1U in control rats and both XBP1U and XBP1S in NBD rat CBLM (A) or HC (B). (C and D) PstI digestion of XBP1 RT-PCR products. Note PstI-insensitive XBP1S band in NBD but not control rat CBLM (C) and HC (D). (E) Sequencing of XBP1U and XBP1S from CBLM and HC confirms excision of the 26-nucleotide intron from XBP1S in NBD rats. (F and G) SYBR Green real-time PCR analysis of total XBP1 mRNA in CBLM and HC of control (n = 5) and NBD (n = 7) rats. Note that XBP1 mRNA expression levels are unchanged in NBD CBLM (F) but are significantly increased in NBD HC (G: 1.3-fold increase in NBD; ANOVA, P = 0.005). (H and I) Western blot analysis of XBP1U (33 kDa) protein levels in CBLM and HC cytoplasmic and nuclear extracts from control (n = 4) and NBD (n = 4) rats. Representative blots are shown. (H) Relative band densities for CBLM extracts revealed a trend toward decreased XBP1U protein in cytoplasmic extracts (2.49-fold decrease in NBD; ANOVA, P = 0.065) and a significant decrease in nuclear extracts (1.72-fold decrease in NBD; ANOVA, P = 0.017) from NBD rats compared to controls. (I) Relative band density for HC extracts revealed significant decreases in XBP1U protein in both cytoplasmic (2.63-fold decrease in NBD; ANOVA, P = 0.033) and nuclear (3.02-fold decrease in NBD; ANOVA, P = 0.018) extracts from NBD rats compared to results for controls. Asterisks indicate P values of <0.05.
XBP1 mRNA levels are also induced during ER stress through an ER stress response element (ERSE) in the XBP1 promoter (77). To detect total XBP1 mRNA, we performed real-time PCR using primers that amplify both unspliced and spliced transcripts of XBP1 mRNA. We did not find differences in total XBP1 mRNA in NBD CBLM (Fig. (Fig.4F),4F), but total XBP1 mRNA was increased in NBD HC (1.3-fold increase; ANOVA, P = 0.005) (Fig. (Fig.4G4G).
We assessed XBP1 protein levels in cytoplasmic and nuclear extracts from control and NBD CBLM and HC in Western blots using antibodies reactive with both constitutively expressed XBP1U (33 kDa) and XBP1S (54 kDa). In NBD CBLM there was a trend toward decreased XBP1U protein in the cytoplasm (2.49-fold decrease; ANOVA, P = 0.065) and a significant decrease in the nucleus (1.72-fold decrease; ANOVA, P = 0.017) (Fig. (Fig.4H).4H). In NBD HC, XBP1U protein was significantly decreased in both the cytoplasm (2.63-fold decrease; ANOVA, P = 0.033) and the nucleus (3.02-fold decrease; ANOVA, P = 0.018) (Fig. (Fig.4I).4I). XBP1S was not detected in any extract.
ATF6 cleavage in NBD rats.
Accumulation of unfolded or misfolded proteins in the ER results in release of ATF6 from Bip and permits ATF6 transport to the Golgi compartment. Cleavage of ATF6 by site-1 and site-2 proteases generates the activated cytosolic form of ATF6 (ATF6p50), which translocates to the nucleus to orchestrate transcriptional activation of genes encoding ER chaperones and folding enzymes (22, 48, 62, 75). We assessed the cleavage status of ATF6 in protein extracts from NBD CBLM and HC by Western blotting (Fig. (Fig.5A).5A). ATF6p50 protein levels were significantly increased in NBD CBLM (Fig. (Fig.5B)5B) (1.33-fold increase; ANOVA, P = 0.0073) and HC (Fig. (Fig.5C)5C) (2.74-fold increase; ANOVA, P = 0.0084). ATF6p50 was not detectable in NBD or control CBLM or HC nuclear extracts (data not shown).
Enhanced cleavage of ATF6 in NBD rats. (A) Western immunoblots of ATF6 cleavage (ATF6p50) in cytoplasmic extracts from PND28 control and NBD CBLM (upper left panel) or HC (upper right panel). Corresponding signals for the GAPDH housekeeping gene are shown (lower panels). (B) ATF6p50 band density, relative to that for GAPDH, from control (n = 4) or NBD (n = 4) rat CBLM was significantly higher for NBD rats than for controls (1.33-fold increase for NBD; ANOVA, P = 0.0073). (C) ATF6p50 band density, relative to that for GAPDH, from control (n = 4) or NBD (n = 4) rat HC was significantly higher for NBD rats than for controls (2.74-fold increase in NBD; ANOVA, P = 0.0084). Asterisks indicate P values of <0.05.
CHOP mRNA and protein expression in NBD rat brain.
The CHOP promoter contains cis-acting elements (ERSEs and amino acid regulatory elements) that regulate transcriptional activation. The downstream UPR effectors, ATF4, XBP1S, and ATF6p50, converge on elements in the CHOP promoter to induce CHOP transcription (49). SYBR Green real-time PCR confirmed increases in CHOP mRNA in PND28 NBD CBLM (Fig. (Fig.6A)6A) (2.77-fold increase; ANOVA, P < 0.0001) and HC (Fig. (Fig.6B)6B) (4.14-fold increase; ANOVA, P < 0.0001).
Increased CHOP mRNA and protein in NBD rats. (A and B) Real-time PCR analysis of CHOP mRNA in CBLM (A) and HC (B) of PND28 control (n = 5) or NBD (n = 7) rats. CHOP mRNA was significantly increased for NBD rats relative to results for control CBLM (A: 2.77-fold increase in NBD; ANOVA, P < 0.0001) or HC (B: 4.14-fold increase in NBD; ANOVA, P < 0.0001). (C and D) Representative Western immunoblots for CHOP protein in cytoplasmic (top panel) or nuclear (middle panel) extracts from control or NBD rat CBLM (C) or HC (D). Corresponding cytoplasmic immunoblot signals for GAPDH are shown (bottom panel). (E and F) Determination of CHOP protein band density in cytoplasmic and nuclear extracts from control (n = 4) or NBD (n = 4) rat CBLM (E) or HC (F). CBLM CHOP protein levels were significantly increased in both cytoplasmic (E: 21.4-fold increase in NBD; ANOVA, P < 0.0001) and nuclear (E: 11.34-fold increase in NBD; ANOVA, P = 0.0003) extracts from NBD rats, relative to results for controls. HC CHOP protein levels were significantly increased in both cytoplasmic (F: 26-kDa band, 3.48-fold increase in NBD; ANOVA, P = 0.043; and 30-kDa band, 7.49-fold increase in NBD; ANOVA, P = 0.009) and nuclear (F: 5.47-fold increase in NBD; ANOVA, P = 0.02) extracts from NBD rats, relative to controls. Asterisks indicate P values of <0.05.
To test whether increased CHOP mRNA correlated with increased CHOP protein levels and to assess distribution in cellular compartments, cytoplasmic and nuclear extracts from NBD and control CBLM and HC were analyzed by Western blotting. CHOP-specific antibodies detected a single, 30-kDa band in cytoplasmic and nuclear extracts of NBD CBLM and in nuclear extracts of NBD HC; no distinct bands were detected in extracts from controls (Fig. 6C and D). In cytoplasmic fractions of NBD HC, an additional 26-kDa band was present (Fig. (Fig.6D)6D) that may represent unphosphorylated CHOP protein (45, 70). Quantitation revealed prominent increases in the 30-kDa protein in both NBD CBLM cytoplasmic (21.4-fold increase; ANOVA, P < 0.0001) and nuclear (11.34-fold increase; ANOVA, P = 0.0003) extracts (Fig. (Fig.6E).6E). Significant increases were also found in NBD HC cytoplasmic extracts for the 26-kDa (3.48-fold increase; ANOVA, P = 0.043) and 30-kDa (7.49-fold increase; ANOVA, P = 0.009) CHOP species and in NBD HC nuclear extracts for the 30-kDa nuclear CHOP species (5.47-fold increase; ANOVA, P = 0.02) (Fig. (Fig.6F6F).
Hippocampus-specific induction of Bip and PDI in PND28 NBD rats.
The ER chaperone, Bip, serves as a key sentinel for ER disturbances, monitoring the folding status of proteins and controlling the activation state of the UPR transducers. Bip is induced at the transcriptional level during ER stress to cope with the accumulation of misfolded proteins in the ER (33). XBP1S and ATF6p50 induce Bip mRNA expression through ERSE elements in the Bip promoter, while ATF4 activates Bip through an ERSE-independent, ATF-CRE-like binding site (42, 76, 78). Thus, Bip induction serves as an indicator of ER stress and UPR induction. Overexpression of Bip reduces ER stress-mediated apoptosis and attenuates CHOP induction (71). Real-time PCR analysis of Bip expression in NBD revealed no alteration in CBLM (Fig. (Fig.7A)7A) but a significant elevation in NBD HC relative to results with control rats (Fig. (Fig.7B)7B) (1.61-fold increase; ANOVA, P = 0.0001).
Hippocampus-specific increases in Bip mRNA and PDI protein levels. (A and B) SYBR green real-time PCR analysis of Bip transcript levels in CBLM (A) and HC (B) of PND28 control (n = 5) or NBD (n = 7) rats. Bip mRNA was not significantly altered in NBD CBLM (A). Note significant increase in HC Bip mRNA in NBD rats relative to controls (B: 1.61-fold increase in NBD; ANOVA, P = 0.0001). (C) Representative Western immunoblot analysis for PDI protein in CBLM and HC of control or NBD rats (upper panels). Corresponding immunoblot signals for the GAPDH housekeeping gene are shown (lower panels). (D and E) Quantitation of PDI immunoblot band density, relative to GAPDH signal, from control (n = 4) or NBD (n = 4) rat CBLM (D: not significant) and HC (E: 1.73-fold increase in NBD; ANOVA, P = 0.0009). Asterisks indicate P values of <0.05.
PDI is induced by ER stress and plays a crucial role in regulating the proper folding of proteins in the ER by catalyzing disulfide bond formation (43). Neuronal overexpression of PDI has also been shown to promote survival in response to stresses that induce ER dysfunction (64). We evaluated protein levels of PDI in extracts from NBD and control rats. PDI-specific antibodies detected the expected 59-kDa species for PDI in both control and NBD CBLM and HC (Fig. (Fig.7C).7C). Although PDI protein levels were similar in CBLM extracts from NBD and controls (Fig. (Fig.7D),7D), PDI levels were increased in NBD HC (Fig. (Fig.7E)7E) (1.73-fold increase; ANOVA, P = 0.0009).
ER stress in PCs.
Previous reports have identified apoptotic neurodegeneration of specific neural populations in NBD, including Purkinje cells (PCs) (25, 72). Calbindin staining of PND28 NBD rat CBLM confirmed PC loss (Fig. (Fig.8A,8A, brackets). H&E staining identified degenerating PCs in NBD, with pyknotic nuclei or signs of nuclear fragmentation (Fig. (Fig.8B).8B). Regional expression of CHOP was assessed in brain sections of PND21, PND28, and PND84 NBD and controls by immunofluorescence microscopy. Specific CHOP staining was not present in control rat CBLM (Fig. (Fig.8C).8C). At PND21, a time point preceding loss of PCs, we did not detect differences in the CHOP signal between NBD and the controls (data not shown). However, at PND28, the CHOP signal was dramatically increased in many PC bodies and dendrites in rats with NBD (Fig. (Fig.8C).8C). To test whether UPR activation localized to PCs in rats with NBD, we assessed eif2α phosphorylation using antibodies specific to P-eif2α. P-eif2α was observed in PCs of rats with NBD but not in controls (Fig. (Fig.8C).8C). We also evaluated Bip and PDI immunofluorescence. There was no detectable difference in PC signal for Bip and PDI between control and NBD rats (Fig. (Fig.8C).8C). Faint PDI fluorescence was detected in a few glial cells in the CBLM of NBD rats (data not shown). Double-label immunofluorescence experiments using antibodies to CHOP and ATF4 indicated colocalization in NBD rat PCs and enhanced ATF4 signal in PND28 NBD PCs relative to results with control rats (Fig. (Fig.8D8D).
ER stress in NBD rat PCs. (A) Calbindin staining in the CBLM of PND28 NBD rats; note gaps in the PC and molecular layers (brackets). (B) H&E-stained PCs from PND28 NBD rat; note degenerating PCs with pyknotic and fragmented nuclei (arrows), with normal PC shown for comparison (arrowhead). (C) Representative CHOP, P-eif2α, Bip, and PDI immunofluorescence of PND28 control (left panels) or NBD (right panels) rat CBLM. Note intense CHOP and P-eif2α immunoreactivity in PCs of NBD rats compared to results for control rats. Bip and PDI signals are equivalent in control and NBD rat PCs. (D) Double-label immunofluorescence analysis of CHOP (red) and ATF4 (green) in PND28 NBD rat CBLM (merged image shows colocalized signal in PCs, arrowheads). ATF4 immunofluorescence in PND28 control rat CBLM shown for comparison (bottom panel). Abbreviations: gcl, granule cell layer; ml, molecular layer. Micron bars, 100 μm.
ER stress in hippocampal astrocytes.
CHOP induction in NBD HC occurs in conjunction with induction of Bip and PDI. Thus, we examined the distribution of each of these proteins by immunofluorescence microscopy. CHOP was confined to astrocytes (Fig. (Fig.9A).9A). CHOP staining was weak in control HC; however, at PND21, PND28, and PND84, NBD HC had marked CHOP signal in astrocytes in the molecular layer of the dentate gyrus, stratum lacunosum moleculare, stratum radiatum, and stratum oriens (Fig. (Fig.9B,9B, representative PND28 rats). Surprisingly, dentate gyrus granule cell neurons, which rapidly degenerate between 3 and 5 weeks postinfection in NBD rats, did not overexpress CHOP at PND21 or PND28. Whereas P-eif2α signal was weak in controls, intense P-eif2α immunoreactivity was observed in astrocytes in NBD HC (Fig. (Fig.9B).9B). Bip fluorescence was weak in controls and localized primarily to neurons. In NBD rats, however, enhanced Bip staining was readily detected in astrocytes in HC. PDI levels were high in neurons in the HC of controls (i.e., dentate gyrus granule cell neurons and pyramidal neurons of CA3-CA1), but astrocytes were only weakly stained (Fig. (Fig.9B).9B). In contrast, PDI signal was increased in numerous astrocytes in the HC of NBD rats (Fig. (Fig.9B9B).
Induction of ER stress in hippocampal astrocytes (A and B) and Chop expression in diverse, degenerating neural populations (C and D) in NBD rats. (A) Double-label immunofluorescence for CHOP (left panel, red) and GFAP (middle panel, green) in HC of NBD rats (right panel, merged image). (B) CHOP, P-eif2α, Bip, and PDI immunofluorescence in dentate gyrus of PND28 control (left panels) or NBD (right panels) rats. Note intense immunoreactivity for CHOP, P-eif2α, Bip, and PDI in NBD rat astrocytes in dentate gyrus molecular layer and hilus compared to controls. (C) Chop immunofluorescence (left panels) and degenerative morphology (arrows in right panels) in cerebellar granule cell neurons (CBLM gcl), cortical neurons (Cortex), hippocampal CA3 pyramidal neurons (CA3), thalamic neurons (Thal.), and hypothalamic neurons (Hypothal.). (D) Chop immunofluorescence in mitral cells in the olfactory bulb of PND28 NBD rat. Note intense nuclear staining (wide arrowheads) and localization to nuclear bodies (narrow arrowheads). (E) Anti-BDV nucleoprotein staining of mitral cells in NBD rats. (F) H&E staining shows degenerating mitral cell (arrow) and normal mitral cell (arrowhead) in NBD rat. Abbreviations: gcl, granule cell layer; ml, molecular layer; Hi, hilus. Micron bars, 100 μm.
CHOP expression in other brain regions.
The CBLM and HC are not the only brain regions reported to be affected in NBD (72); thus, we investigated CHOP expression throughout the brains of NBD rats. Neuronal CHOP expression was not restricted to PCs. Subcellular localization of CHOP varied in individual neurons, including PCs, localizing to the cytoplasm only, the nucleus only, both the cytoplasm and the nucleus, or intensely stained nuclear bodies. Scattered cerebellar granule cell neurons, numerous cortical neurons, rare CA3 pyramidal neurons, and numerous thalamic and hypothalamic neurons with a strong CHOP signal were found in NBD rat brains (Fig. (Fig.9C).9C). We also detected clear signs of neurodegeneration in H&E-stained sections from these same regions (Fig. (Fig.9C;9C; note nuclear pyknosis and fragmentation in neurons [arrows]). CHOP staining was detected in numerous mitral cells in the olfactory bulb of NBD rats (Fig. (Fig.9D).9D). Mitral cells stained intensely with antibodies for BDV nucleoprotein (Fig. (Fig.9E),9E), and some showed classic histological signs of neurodegeneration (Fig. (Fig.9F;9F; note mitral cell with condensed chromatin; normal mitral cell is shown for comparison). While the density of astrocytes expressing CHOP was highest in the HC of NBD rats, CHOP-positive astrocytes were observed in the cortex, striatum, and thalamic and hypothalamic regions (data not shown). Neither neuronal nor astrocytic CHOP expression was detected in corresponding regions of control rats (data not shown) or in regions of NBD rats where histological signs of neurodegeneration were not evident.
DISCUSSION
Viral infections have been shown to induce apoptosis through ER stress. However, the pathological consequences of ER stress associated with viral disease have not been well established, since most reports have focused on in vitro infections. In this study we demonstrate that UPR activation and ER stress are correlated temporally and anatomically with neural damage in NBD. Cell death occurs when ER quality control mechanisms are overwhelmed by excessive or prolonged stress; the balance between apoptotic and quality control UPR signaling likely determines cell fate. Our investigation of the ER stress-induced proapoptotic gene CHOP and the ER quality control chaperones Bip and PDI show cell type and regional discordance. In CBLM, increased expression of CHOP in degenerating PCs was not associated with compensatory increases in Bip and PDI. In contrast, astrocytes, a cell type that survives in NBD, expressed Bip and PDI as well as CHOP. These results suggest that distinct cell types may harbor unique, deficient, or developmentally regulated UPR signals that modulate their response to ER stress in central nervous system infection.
UPR activation in NBD.
Microarray experiments in NBD revealed differential expression of ER stress genes and prompted investigations into the activation state of the three transducers of the UPR: PERK, IRE1, and ATF6. PERK-mediated phosphorylation of eif2α during ER stress inhibits global translation, reducing the burden of newly synthesized proteins in the ER (17). Several viruses have developed mechanisms to circumvent translational shutdown. Infection of fibroblasts with HCMV activates ER stress signals, including PERK, but has only minimal impact on eif2α phosphorylation (26). The hepatitis C virus E2 protein binds activated PERK, preventing eif2α phosphorylation, and the herpes simplex virus type 1 γ34.5 protein stimulates eif2α dephosphorylation, resulting in blockade of P-eif2α-mediated translational attenuation (23, 52). In NBD, infection was associated with eif2α phosphorylation in CBLM and HC but did not result in a detectable reduction in overall protein synthesis. Although it is tempting to speculate that BDV has evolved strategies to circumvent translation attenuation despite eif2α phosphorylation, ER stress is limited to discrete cellular populations in NBD and the sensitivity of our assay may be insufficient to detect protein attenuation in heterogeneous cellular homogenates.
CBLM-specific increases in nuclear ATF4 were detected in NBD, and the ATF4 protein localized to PCs. Consistent with this finding, expression of the Asn-S, Hmox1, and Slc3a2 genes, genes regulated by ATF4 (19), were increased in CBLM microarray experiments. Double-labeling experiments colocalized ATF4 and CHOP in PCs. ER stress-regulated ATF4 translation and CHOP induction are markedly attenuated in PERK null and eif2αS51A cells, suggesting that the PERK signaling arm of the UPR may be dominant over IRE1 and ATF6 (18, 60). ATF4 was not detected in NBD HC astrocytes despite increased P-eif2α immunoreactivity. These results suggest divergent, cell-type- and region-specific regulation of ATF4 translation in NBD.
XBP1S encodes a potent transcriptional activator that induces the expression of ER chaperones and genes involved in the degradation of misfolded proteins (35). XBP1S mRNA was detected in NBD CBLM and HC but not in controls. We were unable to detect XBP1S
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CONTENTS
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UPFRONT 04Editorial
HOT LIST 24 COVER STORY
HOT LIST
Get inspired by 30 Canadian HR professionals who are pushing the envelope on culture, benefits, L&D and a whole lot more PEOPLE
GLOBAL HR LEADER
Say hello to the CPO
06 Head to head
22 FEATURES
PERFORMANCE MANAGEMENT
With employers reinventing all HR processes, HRD chats with one expert about how performance management must evolve
What should HR’s role be in supporting employees’ financial well-being?
08 L&D update
A new study says L&D may be missing the mark with older workers
12 Rewards/benefits update
The inside scoop on Facebook’s newest paid leave policy
14 Statistics
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How digital fluency is closing the gender gap at work
16 News analysis
What role should HR play in protecting employee data?
21 Opinion
38 FEATURES
INSIDE HR
When Commvault undertook a massive global rebrand, Jesper Helt helped to make it happen. He tells HRD how
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Dave Ulrich outlines how HR can add business value in 2017
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PEOPLE 47 Career Path
How Lorne Rubis went from coaching kids to transforming organizations
48 Other life
Kicking it with HR consultant, author and black belt Karen Gately
HubSpot’s Katie Burke explains how the company is making innovative approaches to HR work
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MONEY IN THE BANK
The benefits of supporting employees’ financial well-being are clear. So what’s the best strategy?
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UPFRONT
EDITORIAL www.hrmonline.ca SPRING 2017
CEO, CHRO, CFO … CPO?
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any will be looking ahead to 2017 with relief and perhaps some trepidation. If nothing else, 2016 proved we are, as the saying goes, living in interesting times – if not downright disruptive ones: Brexit, a surprise (for some) election result in the US, military flare-ups in the Middle East, stock market volatility … the list goes on. In terms of HR, 2016 saw further evolution of the once-humble ‘personnel’ function. Historians may look back on the years between 2015 and 2020 as HR’s ‘coming of age,’ whereby the function finally gained the analytical tools and capabilities to help it become a core member of the corporate decision-making team. Looking ahead, some have hypothesized that the executive board of the future may have a new member – the chief performance officer – which will combine HR, finance, planning, operations and strategy. The new role has emerged in the aftermath of the financial crisis, as a result of increasing globalization, and the growing impact of digital disruption in the workplace.
The executive board of the future may have a new member – the chief performance officer – which will combine HR, finance, planning, operations and strategy The CPO title first rose to prominence in 2009, when US President Barack Obama appointed a member of his staff, Anthony Politano, to the position. Politano described the role as having the ‘six Cs’: “Collect, consolidate, and condense performance-related data; communicate the results; collaborate with others; and control and govern the process.” It’s a combination of skills that might have once rested with the CFO or, more recently, the CHRO. The advent of the CPO is a sure sign that, in light of volatile market conditions, the most successful companies will have key functions working together to drive performance across the business.
Iain Hopkins, editor
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UPFRONT
HEAD TO HEAD
Should employers assist with financial well-being? Canadians are stressed about their personal finances, which means stress-related illnesses are on the rise. What role should HR play?
Ben Bazinet
Vice-president of human resources Horizon North “Employers cannot overlook the impact an employee’s financial well-being has on their health, productivity and engagement. Many simple, cost-effective strategies can be deployed to help the employee take control of their economic behaviours. A financial advisor might be willing to share valuable information in exchange for pitching their services to potential new clients. Government agencies can come and speak about specific programs. Hosting these presenters in a lunch-andlearn format ensures they don’t cost valuable productivity hours. The goal should be to give employees the tools necessary to manage their own financial health, not to manage it for them.”
Alec Bashinsky
National partner, people and performance Deloitte “Many organizations provide both internal and external-vendor financial planning services to their employees. In some cases, this is seen as part of an employee benefits offering or as part of a broader well-being strategy. While there are many pros and cons to assisting employees with their financial well-being, my view is this is something an organization should not be involved with apart from providing access to services. If the advice fails or backfires, the employee will blame the organization for approving or endorsing this assistance, and this has the propensity to disengage the employee or create an adversarial position between them and the organization.”
Kim Jones
Lead, talent innovation and staffing ATB Investor Services “Investing in the financial well-being of employees is overlooked by many companies, yet it is essential for any organization that aspires to be world-class. Financially fit and savvy employees lead to higher engagement and productivity. At ATB, we recognize that financial wellness is part of an individual’s total well-being and invest in the ‘personal equity’ of all employees. Besides offering preferred banking rates, outstanding perks and a bank branch for the exclusive use of employees (which might not apply to other industries), we also offer one of the best flexible pension plans, a unique benefits program, financial education and financial literacy workshops.”
FEELING THE FINANCIAL STRAIN Four out of five employers are feeling the consequences of their workers’ stress over financial issues reflected in job performance, according to a report released last year by the International Foundation of Employee Benefit Plans [IFEBP]. “Employers are reporting more financial challenges among employees today than five years ago and are seeing these challenges reflected in the day-to-day operations of their workplace,” said IFEBP director of research Julie Stich. “Nearly half of organizations rate their workforce as only a little bit or not at all financially savvy.” According to this report, workers are most likely to stress over debt, which was a factor for 66% of respondents; saving for retirement, which was a concern for 60% of workers; and saving for or funding children’s education, which just over half of respondents reported as a worry. The IFEPB found that two-thirds of North American employers offer their employees some kind of financial education.
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UPFRONT
L&D UPDATE
Why older workers are missing out on L&D Mature employees often feel pressure to perform well, which leaves them feeling stressed, says a new study
by young coworkers and when they worked in manual occupations. There is, however, a solution – the research found that two types of management practices reduce stereotype threat and increase engagement among mature-age workers. The first type is high-performing practices that focus on training employees, rewarding them for good performance and encouraging them to participate in organizational decisions. The second type is a specific focus on mature workers – updating their skills, redesigning
“Organizations rarely offered the practices that focused directly on mature-age workers”
Well-meaning L&D initiatives can easily miss the mark with older employees, leaving them stressed and disengaged. That’s the warning from a new report, published in the Academy of Management Journal, which compiled information from more than 600 employees aged 45 and older. One its key findings was that many respondents experienced ‘stereotype threat,’ which happens when this age group feels extra pressure to perform well, making work more stressful.
NEWS BRIEFS
Carol T. Kulik, Sanjeewa Perera and Christina Cregan, the trio that led the study, pointed out that this is not only bad for employees, but for organizations, too. “Disengaged workers don’t perform to their full capability and can cost an organization 30% of their salary in lost productivity,” they said. According to the study, mature-age workers experience the most stereotype threat in three situations: when they reported to young managers, when they were surrounded
INSEAD named top MBA program in the world
The Financial Times has named INSEAD’s MBA program as the world’s best for the second year running. To achieve the top ranking, INSEAD scored highly in alumni career progress, international diversity and idea generation. In 2016, INSEAD – which has campuses in France, Singapore and Abu Dhabi – became the first school with a oneyear MBA program to take the top spot. Rounding out the top five for 2017 are the business schools at Stanford, Wharton, Harvard and Cambridge.
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jobs to accommodate their physical needs, giving them opportunities to mentor others or allowing them to ease into retirement. According to the researchers, these agespecific practices send a clear signal to mature-age workers that the organization cares about them and their needs. The two types of practices have independent effects, so the mature-age worker will be most engaged when their organization offers both. “Unfortunately, we found that organizations were more likely to offer the highperformance practices. They rarely offered the practices that focused directly on mature-age workers,” the authors said. “Eventually, [those workers] disengage and feel less involved and enthusiastic about their work.”
Amazon rolls out plan to help underperformers Internet giant Amazon is introducing a new training program called Pivot, designed to help underperforming employees. Pivot pairs struggling learners with ‘career ambassadors’ who provide guidance and support for a fixed period of time to employees “who show a sustained period of underperformance, despite coaching and support.” The push comes as Amazon seeks to improve its unpopular performance improvement plan, which was designed to put pressure on those deemed to be failing to reach their goals.
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28/02/2017 7:04:17 AM
UPFRONT
L&D UPDATE Q&A
Driving growth through L&D George Matti Senior consultant CCM CONSULTANCY
Fast fact CCM Consultancy (ccmconsultancy. com) also represents TMI World and TACK International, two of the oldest learning and management consultancies in the world, in Canada, the US, and the Middle East and North Africa
How has the L&D arena changed over the last five years? While the fundamentals of L&D remain the same, the space has evolved in several ways. We have seen use of gamification on the rise, with learners keen on having more fun in learning. While this is not a new concept, its application in the L&D space is relatively new. Over the last few years, e-learning was on the rise, and many companies thought this was sustainable. The fact is, we continue to see that learners are not as engaged with e-learning – the dropout rate is very high, and the application of the learning is not where companies had hoped it would be. So the ‘happy medium’ has been bite-size learning coupled with online resources and gamified activities using apps and other tools.
How important is emotional intelligence when it comes to leadership development? This is by far one of the biggest game-changers in companies today, and leaders at all levels must have a keen understanding of emotional intelligence and its application in the workplace. This doesn’t mean emotional intelligence replaces cognitive intelligence or experience; it simply means that for leaders to excel today, they must enhance their level of selfawareness, self-motivation and self-management, as well as increase their levels of empathy. Unlike intelligence that is often measured through IQ, our EQ can be enhanced through learning and practice. We have the ability to learn new skills in this area and demonstrate our capability with our team members. EI can make the difference in employees
Destination determines learning success
Whether learners are likely to succeed in online courses largely depends on where they live. A study published in the journal Science found that of the 25 million people who enrolled in massive open online courses [MOOCs] between 2012 and 2015, those in less developed countries completed the courses at a lower rate than those in more developed parts of the world. “Judged by completion rates, MOOCs ... reflect prevailing educational disparities between nations,” said lead study author René Kizilcec.
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staying, even during challenging financial times, or leaving due to a bad leader.
Do you think some employers overlook the importance of lifelong learning in leaders? I believe too many employers overlook this area. There is so much focus on the front-liners that we tend to forget that leaders need to be invested in – their skills improved and updated – to ensure that they lead by example. Lifelong learning should be looked at as a top-down approach, and leaders should be at the forefront of learning and being upskilled. Time and again, studies have proven that satisfied employees have a direct impact on the company’s profitability and growth. This cannot be achieved by solely investing in the front line, but rather requires engaging leaders and creating a sustainable learning plan for them.
If you had one piece of advice for employers who want to improve L&D within their organizations, what would it be? I would suggest that they engage their employees and find out what matters to them. We often think we can figure out the employees’ needs simply by asking the line managers or mapping their capabilities against the required competencies. While this can go a long way and is often more than most are doing, engaging our staff will give them a sense of empowerment and inclusion. The ideas that will come from the team will surprise most, and you can enhance participation during the process.
Will religious training be the next L&D trend?
One of the world’s largest advisory firms has launched a specialized training program that seeks to improve religious understanding within organizations all over the world. The program, Religious Literacy for Organizations, was designed by EY to drive inclusivity and show the positive impact it has on business process and performance. It’s estimated that approximately 84% of the world’s population identifies as religiously affiliated, yet religion remains a highly sensitive subject within many businesses.
Gates says tax on robotic workers should fund L&D
Robotic workers shouldn’t be exempt from paying income tax, according to Microsoft co-founder Bill Gates, who says the funds could then be used to retrain employees for jobs that will never be threatened by automation, such as teaching, eldercare or special needs services.“All of those are things where human empathy and understanding are still very unique, and we still deal with an immense shortage of people to help out there,” Gates said in an interview with digital news outlet Quartz.
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2017-02-22 28/02/2017 7:04:314:19 AMPM
UPFRONT
REWARDS/BENEFITS UPDATE NEWS BRIEFS Marshmallows teach HR about delayed gratification
Marshmallows have been involved in a series of studies that examined the link between delayed gratification and life outcomes. The results have proved to be a great teaching tool for managers and employees by demonstrating that patience – and the ability to hold out for a larger reward down the line – can be a key to lasting success. The studies involved children who were given a choice of one marshmallow, provided immediately, or two marshmallows if they could wait a short period of time. The results found that children with the ability to delay gratification tended to have better outcomes later in life.
Career-pathing talks should focus on rewards When asked about the most dissatisfying attributes of their previous jobs, those surveyed in a recent CEB study cited people management (47%), future career opportunities (44%) and recognition (37%). Aaron McEwan, HR advisory leader at CEB, suggested that employers should focus on tackling these key drivers of attrition. The report also advised managers to prioritize informal career-pathing discussions in order to rebuild levels of trust and get staff motivated. While conducting these career conversations, senior staff members should acknowledge the strengths and positive contributions of employees.
Manulife boosts employee benefits package Financial giant Manulife has confirmed an increase in its staff benefits at the start of 2017, providing its entire Canadian workforce with up to $10,000 in coverage for mental health issues.
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Under the new package, all 13,000 Manulife employees and their families will be eligible for annual mental health benefits. “We’re seeking to overtly address the financial barrier [to seeking mental health treatment] with our benefits, but in some ways we are hoping to address the stigma by showing our employees that we care about their mental health,” said EVP of HR Stephani Kingsmill.
R&R firm launches $6.5 million charity fund
International HR service provider Reward Gateway has announced the launch of a $6.5 million fund to “make the world a better place to work.” The RG Foundation will award grants quarterly to charity, nonprofit and community organizations that are helping people improve working conditions and pay. The first beneficiary is the TGG Foundation, a charitable initiative based in Kerala, India, which has received US$50,000. The grant will allow the TGG Foundation to begin work on a Rural Empowerment Hub to provide local residents with jobs, skills development, fairer levels of pay and better conditions.
Employees stressed by bonus schemes Bonus schemes designed to motivate employees may actually end up harming their well-being, according to new research from the University of East Anglia. The study showed that financial rewards tied to performance often resulted in more intense work for employees, leading to work-related stress or poor well-being. “The key thing for managers is to ensure some balance between employee job demands and measurement of rewards offered,” said lead researcher Dr. Chidiebere Ogbonnaya.
Facebook introduces new paid leave The tech giant breaks new ground with a leave policy that will apply to more than just new parents Social media giant Facebook has confirmed a generous new paid leave policy, which will have a significant impact on grieving employees, as well as those who have to care for a sick family member. In early February, COO Sheryl Sandberg announced that the company would be giving all of its employees six weeks of annual leave to care for ailing relatives – but that’s not all. “Starting today, Facebook employees will have up to 20 days of paid leave to grieve an immediate family member [and] up to 10 days to grieve an extended family member,” Sandberg confirmed. “This is personal for me,” she added at Makers Conference, a women’s leadership event held in California. “I lost my husband very suddenly – Facebook provided leave and flexibility, and now we’re doing more.” Sandberg became a widow in 2015 when her husband, fellow tech figure Dave Goldberg, died suddenly of cardiac arrhythmia while on vacation. She returned to work 10 days after his death. Paid family sick time was also added to Facebook’s list of impressive benefits – employees now entitled to take three days off work to care for a family member with a short-term illness, such as a child with the flu. “People should be able both to work and be there for their families,” Sandberg said. “No one
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should face this trade-off. We need public policies that make it easier for people to care for their children and aging parents, and for families to mourn and heal after loss.” The benefits bump is just the latest employee experience enhancement from Facebook, which last year increased its paid parental leave to four months for both men and women. Keen to show the policy wasn’t just an empty offering, founder and CEO Mark Zuckerberg set an early example by taking two months of paternity leave when his daughter, Max, was born. “Studies show that when working parents take time to be with their newborns,
“People should be able both to work and be there for their families. No one should face this trade-off ” outcomes are better for the children and families,” he wrote on his Facebook page after confirming the time off. In Canada, ‘leave’ is classified as the period of time new parents are entitled to take off without pay, with a guarantee they can return to the same job once that leave period ends. Length varies from province to province, as do the eligibility requirements. Maternity leave for biological mothers varies from 15 to 18 weeks. Parental leave, available to both parents, varies from 35 to 52 weeks. Although adoptive mothers are not entitled to maternity leave, adoptive parents are usually entitled to the same parental leave as biological parents.
Q&A
David Brennan General manager ACHIEVERS
Fast fact Achievers is consistently rated one of Canada’s best employers by the Great Place to Work Institute. In 2015, Achievers was ranked 12th out of 50 for Best Medium Workplace and 14th out of 50 for Best Workplace for Women
What HR can expect from R&R What are the major trends you’re seeing in the R&R space right now? Rewards & recognition is no longer a ‘nice to have,’ but has become an essential component of leading organizations’ broader engagement strategies. We’re also finding that a number of companies are moving away from the traditional concept of ‘rewards & recognition’ and toward stand-alone social recognition programs untethered from a rewards component, where recognition is the primary driver of desired behaviours rather than the promise of monetary or other material rewards. We are also seeing some very interesting developments with regards to people analytics and how some of our most forwardthinking customers are using recognition data to inform important HR decisions and give them incredibly granular employee performance and engagement data and insights.
How do you predict the field will change in the next five to 10 years? You’re going to see rewards & recognition programs become totally commonplace in all truly successful companies. But beyond that, R&R will be included as part of a more holistic employee experience solution that will provide employers with insights on how to engage and motivate employees down to an individual contributor level.
What risks do employers face if they fail to adapt with the rapidly evolving industry? Employers risk being left behind. They risk being outperformed on a number of levels and in a number of key business areas where we know R&R has an impact: retention, engagement, customer satisfaction, revenue growth, overall performance. Those are some pretty significant areas of impact. So if you are a business leader and you’re getting left behind in those areas, I would suggest that you might want to give R&R another look. Beyond that, organizations risk being seen as old-fashioned and having their employer brand be negatively impacted as a result. In the current labour market, as defined by an ongoing war for talent, especially among millennials, and by the increasing transparency brought about by social networks like LinkedIn and Glassdoor, your employer brand is more important than ever.
If you had one piece of advice to offer employers in regards to R&R, what would it be? If you’re not already doing it, the time is now to begin thinking about how to link your rewards & recognition program to your company’s key strategic goals. If you’re not using rewards & recognition to move the needle in those areas that are most important to your business, you’re not getting the kind of ROI you should. If delivering a great customer experience is most important to your business, design rewards & recognition programs that tie to key metrics like NPS and CSAT. If increased sales are what you’re after, design a strategic recognition program to incentivize your sales team.
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UPFRONT
STATISTICS
Closing the gender gap
Canada
7/5
71%
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Percentage of respondents who agree that “the digital world will empower our daughters”
Number of countries (out of 31) in which women have attained higher levels of education than men
1/3
76%
55
55
A WORLD OF OPPORTUNITY
using will provide their desired outcome. The work flexibility afforded by newer forms of technology seems to have an outsized effect on women: almost six out of 10 women surveyed who were not currently engaged in formal employment said that working from home part or all of the time would help them find work. A similar proportion reported that having more flexible hours would assist them in their search for employment.
Percentage of men who use digital channels frequently (versus 72% of women)
52
United States
Digital fluency has the potential to narrow the economic divide between men and women WHILE MEN currently have an employment advantage, that gap is closing, due in large part to a significant uptick in outcomes for women driven by digital fluency. According to a recent Accenture study, nations with higher rates of female digital fluency also have higher overall rates of gender equality. A digitally fluent person can decide when to use specific technologies to achieve their desired outcome, and also articulate why those tools they are
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Accenture defines digital fluency as the combined result of factors including education, employment and advancement at work. The firm rated each country it studied on a scale of 100 points on these core components to produce an overall score, as well as a separate digital fluency rating that covers digital use only. The US, Netherlands and Australia took the top three spots overall, while Canada came in seventh. However, Canada ranked fifth in terms of its digital fluency score. Overall rank/digital fluency rank
61%
Overall score
Percentage of women in developing countries who want to start a new business in the next five years
Digital fluency score
Source: “Getting to Equal,” Accenture, 2016
LARGEST GAPS
ACCELERATING THE PACE
Of the five countries with the largest gap in digital fluency based on gender, the top two are both in Asia; the remaining three countries are in continental Europe.
In a scenario in which women were enabled to become digitally fluent at double the speed, gender equality could be reached decades earlier than the current date forecast of 2065 for developed countries and 2100 for the developing world.
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60
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2015
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2X SPEED: 45 years to reach gender equality
10 0 Japan
Singapore
Women
France
Switzerland Netherlands
Developing countries
Men Source: “Getting to Equal,” Accenture, 2016
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Developed countries
Source: “Getting to Equal,” Accenture, 2016
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Netherlands
2/1
Nordic
52
63
4/2
51
58
Switzerland United Kingdom
5/4
49
45
9/10
53
48 Germany
Austria
14/16
Ireland
45
8/12
40
15/11
52
63
47
United Arab Emirates
44
11/6
52
France
Spain
6/7
47
46
52
10/13
44
46
Singapore
12/15 Argentina
13/19
41
42
42
Australia
35
3/8
52
50 Source: “Getting to Equal,” Accenture, 2016
REACHING AN AGREEMENT
SMALLEST GAPS
An overwhelming majority of respondents agreed that women’s employment opportunities increase as their digital fluency rises, with only a minor discrepancy between the genders.
The countries with the smallest gender gaps tend to be developed nations – three of the top four are located in Europe. In three of the five countries, women’s digital fluency outstrips that of men.
RESPONDENTS WHO BELIEVE WOMEN’S EMPLOYMENT OPPORTUNITIES INCREASE AS DIGITAL FLUENCY RISES
60 54 50 53
52
49 48
47 47
50
40
72% Women
68% Men
35
30
32
20 10 0 UK
Ireland
South Korea
Women Source: “Getting to Equal,” Accenture, 2016
Spain
Argentina
Men Source: “Getting to Equal,” Accenture, 2016
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UPFRONT
NEWS ANALYSIS
Data breaches and HR Several high-profile cases around the world have highlighted the need for companies to prepare for cyberattacks. What role can HR play? THE NEW cybersecurity buzzword ‘insider threat’ is a nod to the growing understanding that internal data security deserves as much – if not more – attention than the external threats we know and fear. “While high-profile data breaches through external hacks are often those that make the news, the reality is that the majority of data breaches fly under the radar,” says Guy Eilon, senior director and general manager of Forcepoint, a data security advisory firm. Indeed, around 53 million data security incidents took place last year alone, according to a global report by IBM. “While the headlines would lead us to believe that
to a phishing scam) or a purposeful leakage (when an employee has a specific motive to steal sensitive data). To provide just one example, in late 2016, global recruiting firm Michael Page had its security breached in an accidental leak. The records of 711,000 job candidates from China, the UK and the Netherlands were accessed by two individuals “without malicious or fraudulent intent,” according to the firm. Michael Page reported that the leak occurred on a server used by its third-party IT service provider, Capgemini. In order to combat the threat of employees stealing information, HR must
“While high-profile data breaches make the news, the reality is that the majority of data breaches fly under the radar” Guy Eilon, Forcepoint these are due to sophisticated cybercrime from foreign adversaries, [many] of these incidents were in fact due to the company’s biggest asset: their employees,” Eilon says. This type of ‘insider threat’ encompasses a number of potential dangers that stem from internal vulnerabilities, including accidental leakage (when an innocent employee leaves sensitive data lying around), social engineering (when an employee falls victim
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work closely with the IT team, says Lynne Lewis, intellectual property partner at law firm Minter Ellison. In particular, Lewis says, it’s important to ensure that there are comprehensive systems in place so that employees engaging in potentially harmful behaviour can be tracked. Employers have the option of accessing technology that allows them to see if an employee has been downloading onto
USB keys, or if they have been taking their computer home, hooking it up to devices and downloading or printing from it. However, Lewis has come across organizations, quite often large ones, that don’t have sound IT systems in place – for example, there may not be any logs kept of USB keys that are plugged in and out of workplace computers. “If I’m running a case, I want to be able to show that the night before someone actually departed or the night before they handed in their resignation, a USB key was inserted into their work computer and that XYZ was copied over onto that,” Lewis says. “HR must work with the IT team to get good tracking and lockdown technology in place ahead of time.” HR teams also need exceptional internal communication in place whenever an employee resigns, she adds. “I’ve seen many circumstances where the information doesn’t necessarily get told
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LARGE-SCALE INTERNAL LEAKS AROUND THE WORLD RED CROSS BLOOD SERVICE (2016) Personal data of 550,000 blood donors – including names, gender, address and date of birth, as well as information on ‘at-risk sexual behaviour’ – was accidentally published to the public. VODAFONE (2013) Using privileged insider access, an IT contractor working for Vodafone copied the names and bank account details of 2 million customers from a server located in Germany. MOSSACK FONSECA (PANAMA PAPERS, 2016) Sensitive data on clients, including high-profile politicians, criminals and professional athletes, was leaked from law firm Mossack Fonseca. The roughly 11.5 million documents included emails, contracts, scanned documents and transcripts.
to the correct people, who might notice that something unusual is happening with people’s access to the premises or documents during that period of time,” Lewis says. “Had
in place an exemption to that if they’ve got a basis for it in their system.” It’s critical for HR professionals to develop an understanding that data security is an
“HR must work with the IT team to get good tracking and lockdown technology in place ahead of time” Lynne Lewis, Minter Ellison they known that person had resigned, then they might have actually spoken up about seeing some strange things happening.” It’s also important to be careful with document destruction policies, because sometimes important information can be lost, Lewis says. She says one of her clients introduced a policy whereby everything disappears after 30 days. “That can be really hard to recover,” she says. “But they can put
issue for more than just the IT department, the CIO or the CSO, Eilon says; it impacts recruitment and training, and should involve active participation from every staff member. Eilon adds that staff training can help reduce the instances of accidental and socially engineered data breaches. Moreover, training employees on safe data practices is a great way to improve overall security. He says staff should learn to recognize and report
YAHOO (2016) More than 150,000 US government and military employees were among the victims of the Yahoo data breach, which put their names, passwords, telephone numbers, security questions, birthdates and backup email addresses in the hands of cybercriminals. Experts warned the leak could allow foreign intelligence services to identify employees and hack their personal and work accounts, posing a threat to national security. a phishing scheme email, understand the importance of separating work and personal technology use (including the dangers of third-party apps), and of course, take seriously the importance of having unique and complex passwords across various platforms. Beyond training, however, Elion points out that there are technology solutions that are also an essential part of an effective data protection program. “New-to-market tools are capable of using computer learning to recognize abnormal employee behaviour, identify any potential threats and record those actions to produce a court-admissible piece of evidence, in the form of a playback video, for prosecuting illegal and malicious actions.”
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PEOPLE
GLOBAL HR LEADER
Rewriting the HR rulebook HubSpot is shaking up traditional notions of HR – and it has the IPO success and innovative perks to prove its approach works. HRD sat down with HubSpot’s global VP of culture and experience, Katie Burke, to find out how THE WORD ‘unconventional’ applies not only to HubSpot’s approach to people management, but also to the career of its global VP of culture and experience, Katie Burke. Prior to HubSpot, Burke was the director of marketing and corporate partnerships at Athletes’ Performance and Core Performance (now known as EXOS). This unique mix of communications and marketing skills, combined with HR expertise, has positioned Burke well to tackle her current employer’s challenges.
HR and IPOs Established just over a decade ago, HubSpot develops and markets software products for inbound marketing, including tools for social media marketing, content management, web analytics and search engine optimization. In short, the company knows something about marketing. It’s perhaps the perfect match for Burke, who was handed the responsibility of running the company’s internal and external communications for its IPO early in her tenure. In October 2014, the Boston-based company launched on the New York Stock Exchange with an IPO listing of US$125 million in fresh capital, leaving the company with a market valuation of US$880 million.
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“There’s a very traditional recipe going into an IPO that people want you to follow,” Burke says. “You’re working with your bankers and your lawyers, and you say, ‘OK, here are the financials, here are our growth numbers, here is our projected revenue, and so on.’ We had all of that. Then we kept adding in flags about our culture: our employees, our talent brand,
“It’s not just employees who care about the employer brand; it’s your shareholders, the public market and potential candidates down the line,” says Burke, who adds that HubSpot was wary of falling into the trap of other similarly sized businesses, which have gone down the IPO path while claiming that culture was critically important, only to fail dismally.
“It’s not just employees who care about the employer brand; it’s your shareholders, the public market and potential candidates” our community and the net worth effect all this was creating.” Part of the reason for this focus, she adds, was due to the number of investors who asked about and deeply cared about the company’s commitment to culture. “They told our founders how a culture of innovation is really what distinguishes outstanding companies from average companies in terms of overall performance. That’s why we knew we had to get the culture piece right.” Indeed, Gallup reports that organizations with above-average levels of employee engagement reap 147% higher earnings per share.
Another of Burke’s key takeaways from the IPO experience occurred on the morning of the IPO. “You have this amazing morning – you walk in [to the stock market floor], and your banner is up, and everyone is excited. You feel energized. And at 11 a.m., they walk you out, they say congratulations, and then another company does their IPO. It’s a very visceral reminder that the world goes on – your party is not everlasting, and you can’t take anything for granted. I think you have to say your IPO is the beginning of your journey as a company, rather than the end. You have to show that to employees.”
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PROFILE Name: Katie Burke Company: HubSpot Title: Global VP of culture & experience HR experience: Two years Previous roles: »»Director, media and analyst relations, HubSpot »»Director of marketing and corporate partnerships, Athletes’ Performance and Core Performance »»Brand manager, EXOS »»Director, Glover Park Group »»Researcher, Booz Allen Hamilton First HR role: Director, talent and culture, HubSpot Qualifications: MBA, Sloan School of Management, Massachusetts Institute of Technology
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PEOPLE
GLOBAL HR LEADER Transparency and autonomy Today, HubSpot employs more than 1,500 people in six locations around the world, including its headquarters in Cambridge, Massachusetts, as well as New Hampshire, Sydney, Dublin, Singapore and Tokyo. A new office will open in Berlin in 2017. The ‘unconventional’ nature of HubSpot’s approach to HR comes down to its two underlying principles: transparency and autonomy. On the former, Burke says that anyone who works at HubSpot, from intern to executive, has access to all of the company’s documents. For example, all staff have access to management team meeting minutes, board meeting outcomes and even the random
who has wanted to be micromanaged; instead, they want latitude to do great work on their own with the support of management. “We set our company goals and directives, but we don’t give people an exact map on how to get there,” she says. While some business leaders might cringe at the thought of training people to think like entrepreneurs, Burke says this mindset is actually helpful to HubSpot. “You feel more accountable to other divisions and to the company. You don’t get that feeling if you’re sitting in the corner with your head down.” If and when ‘HubSpotters’ do leave, it’s not unusual to find the company’s founders and executives acting as angel investors for the
“I think the next few years will see a convergence between HR, marketing and IT. I’m excited by what those changes will bring” musings of the CEO answering off-the-cuff questions from employees. Burke says that being “radically transparent” allows the company to do two things. “One, it prevents people from [problem]solving for themselves or for their team,” she says. “Having to share things and having a bias to share information ensures people are truly ‘solving’ for the company. Second, we believe people do better when they know better. For example, if you’re working on a data project for the data team, and you’re working on something for the marketing team, the bias to share everything prevents people from putting their guard up and saying, ‘That’s my project or my data.’ It encourages collaboration.” She adds that the company’s “super talented” employees likely have career aspirations beyond HubSpot – they may want to start their own company or become a CEO someday. “The idea of learning from other parts of the business just by checking with our company Wiki, for example, is really important for us.” As for autonomy, Burke says she’s never met a talented professional in any division
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new ventures. An active alumni program also gives a boost to these startup companies.
Borrowing from the best How can traditionally risk-averse HR functions learn from companies such as HubSpot, which appears to be rewriting the HR rulebook? For Burke, who says that often the best opportunities in business emerge from a willingness to think differently, there are two initial steps. For HR professionals within large organizations who don’t know where to start and are unsure of what the appetite for change is, she suggests simply asking people what they think. “Don’t ask via a lengthy survey,” she says. “Ask, on a scale of one to 10, how likely you’d be to recommend your company to a friend or colleague. You’ll be surprised by what you get back.” As part of this process, it’s critical to also ask what the company can do better. “I’m constantly amazed at the number of companies who are unwilling to take that step,” Burke says. “Own the accountability coming out of it and ensure you’ll actually take action on it.”
THREE INNOVATIVE STAFF PERKS AT HUBSPOT 1. Unlimited vacation policy. “Unlimited vacations get a lot of attention for the days on versus the days off, but from my perspective, it’s about other things, such as working parents who can do their work without asking for exceptions. Or if you’re someone training for a marathon, you can build your work schedule around what you’re personally passionate about without having to ask for permission.” 2. Free books: “For the last eight years, the company has had a program where you can request a business book. I’m a lifelong learner and avid reader myself, so I love it. Our employees are busy; they have a lot going on. Any friction you create between their ability to learn more and actually doing it can prevent them from taking that leap. So we make it easy for people to learn on their own and learn from other people.” 3. Mystery Dinners: “Three times a year, groups of eight to 10 people sign up for a Mystery Dinner. They have no idea what restaurant or what group of people will be attending until 4 p.m. that day. What we’ve found is that evening of spending quality time over a nice meal goes a long way toward our employees building meaningful connections with people on other teams.” The second step is to ask people to write down what matters most to them about work. Burke often finds that leaders assume that people know what a culture stands for because they have company mission statements or values. But if your newest employee can’t recite what it means or how they feel about it, you’ve failed. Finally, Burke maintains that a culture like HubSpot’s depends on trust. “You can’t fake trust,” she says. “For example, companies that try to launch unlimited vacation policies without a true commitment to autonomy always fail because it’s not about a policy; it’s about the dynamic of trusting employees to do their best at work.”
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UPFRONT
OPINION
GOT AN OPINION THAT COUNTS? Email editor@hrmonline.ca
A great time to be in HR A combination of forces is shifting HR to centre stage, and it’s how HR will handle these forces that will deliver business value, writes Dave Ulrich WE SEE four forces making HR more central to business success: the context of business (STEPED: social, technological, economic, political, environmental and demographic changes), the increased pace of change (VUCA: volatility, uncertainty, complexity, ambiguity), the demise of employee well-being (individuation, isolation, indifference, intensity) and the requirement to be outside in (attend to customers, investors and communities). Collectively, these four forces shift HR to centre stage. In our new book, Victory Through Organization, the first sentence is, “HR is not about HR.” Rather, it’s about attending to these four forces to deliver business value. While it is a great time to be in HR, it is not without downsides. I find three models helpful for HR professionals to respond to the new opportunities.
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Resource demand. The demands on
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Personal resources. HR profes-
HR are increasing through the four forces noted above. HR professionals are being asked to do more. Some of these demands can be moderated (for example, don’t do non-value-added work). But the focus should be on resources. What resources can HR professionals access to help them cope with their increased demands? I see two categories of resources: personal and departmental.
sionals can better respond to increased opportunities when they have the competencies to succeed and a more personalized career map.
Competencies are the skills HR professionals bring to their job. We have spent 30 years studying which HR competencies have personal and business impact. Through our research, we identified nine competencies for HR professionals. Three were foundational (compliance manager, analytics designer and implementer, and technology and media integrator). Three were strategic enablers (culture and change champion, human capital curator, and total rewards steward). But three had the most impact on HR profes-
leader. Within this mosaic, HR professionals may choose to stay within one quadrant (e.g. functional excellence in rewards, learning or organization design), or they may move across quadrants throughout their career (e.g. moving back and forth from centres of expertise to business). When HR professionals proactively define their careers, they can map how they go forward.
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Department resources. HR profes-
sionals seldom succeed by themselves. In our research, we found that the quality of the HR department had four times the impact on business performance as the quality of individual HR professionals (hence the title of our book). Part of creating an effective department is making sure that roles are clear, which is about decision rights, accountabilities and responsibilities. HR departments should be organized to mirror the business organization. If the business has more centralized governance, HR should be more centralized, with common HR practices across the enterprise. But we have come to appreciate that relationships matter as much as (or more than)
“While it is a great time to be in HR, it is not without downsides” sionals around the world: • Being a credible activist allows the HR professional to be invited to the business dialogue and be personally effective. • Being a strategic positioner helps the HR professional deliver the most value to business customers and investors outside the organization. • Being a paradox navigator is the compe tency most connected to delivering business results. So, while HR professionals need to recognize all nine competencies, they especially need to master the three core competencies. With these competencies, HR professionals can take charge of their careers. HR professionals can work in four areas: functional expertise, geographic area, business unit or outside HR. They can work at three stages: individual contributor, manager or
roles. If the structure is mostly right, HR leaders need to build positive relationships throughout the HR community. There have been many studies of positive relationships by psychologists and marriage therapists, which include sharing a common purpose, respecting differences, having realistic expectations, caring for each other, sharing experiences and growing together. As HR moves to centre stage to deliver real business outcomes, the demands need not be threats, but rather opportunities if personal and department resources exist. Dave Ulrich is a university professor, author, speaker, and management coach and consultant. Ulrich is a professor of business at the Ross School of Business at the University of Michigan and co-founder of The RBL Group. For more, visit daveulrich.com.
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LEARNING & DEVELOPMENT INSIGHT
PERFORMANCE MANAGEMENT
Performance management for a new era HRD sat down with performance expert Ian Cullwick to talk about what makes a successful performance management model in 2017 MODERN EMPLOYEES expect their employers to be just that: modern. And those companies that fail to adapt to market forces and expectations of the current and future workforces run the risk of not being able to compete for top talent. In their attempts to modernize and keep up with trends set in motion by companies like Google, Amazon and Facebook, organizations across Canada are finding new ways to conduct core business and HR functions. Performance management is one such HR process that is undergoing fundamental changes at many organizations across the country.
Time for change Over the past year, countless high-profile companies have scrapped traditional performance management processes. Out went performance ratings and formal sit-down conversations; in came more timely and transparent processes – including manager/ employee ‘check-ins.’ In November 2016, Amazon announced it was updating its employee review process and moving away from a model that involved a ‘stack-ranking system’ in which staff members were rated against one another rather than by how well they were meeting their job requirements. “We’re launching a new annual review
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process next year that is radically simplified and focuses on our employees’ strengths, not the absence of weaknesses,” Amazon spokesperson Teal Pennebaker said at the time. “We will continue to iterate and build on the program based on what we learn from our employees.” The reality is that in order to be a successful and mutually beneficial process, performance management has had to become a top
Before being in a position to implement an effective performance management program, an organization must ask itself one simple yet integral question: What do we want to use performance management for? “I would argue that, in and of itself, performance management can have no value,” Cullwick says. “It comes down to contextually understanding the nature of your business and industry sector, and then figuring out how it’s going to add value to your business and employees.” There are some key questions that any organization should carefully consider before designing and implementing a performance management model: • Is it going to enable accountabilities to be strengthened and quality management to be improved? • Is it going to enable team and individual variable pay or bonus determination? • Is it going to enable and support career planning and development? Many organizations use performance management as a way to get a sense of individual and collective interests in terms of career aspirations and progression. However, effective performance management programs
“Performance management is increasingly being recognized as a core management practice, not simply an HR process” Ian Cullwick, Mercer Canada management priority. “In this day and age, performance management is increasingly being recognized as a core management practice, not simply an HR process,” explains Ian Cullwick, a partner in Mercer’s Ottawa office. “That’s largely because it’s the one core management philosophy and practice that, when carried out effectively, can link strategic business direction and priorities to the team and individual employees who will ultimately execute on that strategy and make or break its success.”
can also be used to drive training and development, needs analysis, and related investments in people development and training. There are a number of different ways to apply performance management, so to achieve success, organizational leaders need clear goals that are aligned with the context of their business objectives and needs.
The risks of getting it wrong Although rushing to develop a new performance management model is not a recipe for a
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success, completely ignoring its benefits is sure to have negative organizational consequences. “The reality is that many performance management programs and processes are very unsophisticated or, in some cases, nonexistent,” Cullwick says. “Those with minimal experience who suddenly decide they want to become ‘high-performance driven’ often create ineffective models. Trying to be too sophisticated too quickly can hinder culture and management practices, and cause quite a bit of distraction and misalignment.” Badly designed performance management programs also run the risk of creating false expectations amongst the workforce. If an organization announces the launch of a new performance management system that’s going to drive team and individual bonuses, but the planning and measurement aspects don’t function effectively, leaders will not be in a position to deliver reasonable bonuses that are internally equitable and performance-based. The result is an organization that invests a lot of effort and time, yet receives very little value. Despite a growing awareness around the
importance of providing feedback on performance and setting tangible goals, performance management remains misunderstood by many organizations and industries, particularly by those who do not recognize its value. These organizations are missing opportunities to review, communicate with and incentivize their staff through modern performance management, which can take many shapes and forms. “With technology, data analytics and the information management systems that exist, we’re becoming much more sophisticated with how we plan and measure performance,” Cullwick says. “When well designed and used properly, it can be a major game-changer and value-adder for all types of organizations. Performance management can be a driver that makes a real difference.”
A helping hand To help Canadian firms design and implement effective and optimized performance management models, Cullwick has teamed up with Queen’s University’s Industrial Relations
Centre [IRC] to develop a two-day performance management program, which runs from March 21–22 in Toronto and March 27–28 in Victoria. Cullwick hopes to empower attendees with the knowledge that a robust performance management model must be forward-looking and fundamentally based on the unique contextual needs of an organization’s strategic direction. This program is designed to give participants insights and competency in the key strategies, methods and processes that can be used to realize the benefits of meaningful performance management and successfully drive better accountability, productivity and development. “The objective is to talk about trends and contemporary choices that are available and to help participants understand their own contextual circumstances,” Cullwick says. “We want to help participants better understand the foundational ingredients of performance management: the core basics around different types of planning and measurement approaches and practices, and how different measurement can be used to better focus performance.”
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FEATURES
COVER STORY: HOT LIST 2017
HOT LIST 2017 HRD’s annual Hot List brings together the best of the best in the industry – an impressive group of 30 high achievers in the HR profession
IT’S ALWAYS a pleasure browsing through the final list of HR professionals featured in HRD’s annual ‘who’s who’ guide. These are the people who continue to add value to the profession and increase its standing in the business world. Some of them have appeared on our previous Hot Lists, proving their staying power; others have been featured in the magazine over the past 12 months. Then there are new contenders, the ones who have launched innovative initiatives, tackled tough business-critical challenges or moved the dial on persistent issues like diversity & inclusion. Many have already been acknowledged as industry leaders – they’ve picked up personal accolades from the likes of the Canadian HR Awards, or the employers they work for have been applauded as an employer of choice by organizations such as Aon Hewitt or the Great Place to Work Institute. It’s not all about bright optimism, though; some of the HR professionals on this year’s list are in declining markets or at companies that are struggling to reinvent themselves. But rain, hail or shine, HR continues to prove its worth, as the leaders on the following pages prove.
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HOT LIST INDEX COMPANY
PAGE NAME
COMPANY
PAGE NAME
AG Hair
34
Michele Matthews
Island Savings
28
Ilka Bene
Alberta Health Services
31
Todd Gilchrist
Jones DesLauriers Insurance Management
35
Kristin Coulombe
Aldo Group
33
Maryse Ward
KPMG Canada
30
Linda Speedy
ATB Financial
33
Lorne Rubis
Libro Credit Union
26
Cavell Fraser
Best Buy Canada
28
Kamy Scarlett
Mount Royal University
33
Melanie Peacock
BlueShore Financial
32
Marni Johnson
Nando's Canada
28
Heather Wyllie
Centre for Addiction and Mental Health
34
Paula Harrington
PCL Construction
34
Mike Olsson
City of Vernon
36
Raeleen Manjak
Pelmorex Media
26
Anna Petosa
Coast Hotels
36
Pia Collins
Sierra Systems
36
Stephanie Hollingshead
Colliers International
30
Keri Fraser
SocialHRCamp
28
Jeff Waldman
Enercare
27
Colleen Bailey Moffitt
Spin VFX
30
Leah Jaikaran
Freshii
26
Ashley Dalziel
Traction on Demand
33
Manu Varma
GoodLife Fitness Clubs
26
Alana Free
Ultimate Software
36
Vivan Maza
Greenwin
29
Jenny Affe
Vector Aerospace
34
Monica Rye
HomEquity Bank
30
Lori Sone-Cooper
Worldlynx Wireless
27
Elaine Cruise Smith
“
Stephen has been absolutely delightful and such a positive addition to our team and overall morale .” - Michelle Dulmadge, Director, AltaGas Ltd.
Inclusive Hiring Works Learn more at readywillingable.ca or call 709.437.1504
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FEATURES
COVER STORY: HOT LIST 2017 ALANA FREE
ASHLEY DALZIEL
Vice-president, people and culture
Vice-president, people and culture
GOODLIFE FITNESS CLUBS
FRESHII
In her two decades with GoodLife Fitness, Alana Free has worked her way from receptionist to club manager and eventually L&D manager. She has been responsible for opening new club locations across Canada; now, as VP of people, she oversees a team of 14,000 associates. Over the past year, Free and her team have implemented a new technology system that allows GoodLife to communicate more effectively with its 1.25 million club members while significantly reducing the use of paper. Free also handled the people management side of a significant corporate upheaval, which included providing job transition services such as career counselling, resume-building seminars, education allowance programs and even time off for job interviews while employees still worked with the company.
Amidst concerns about the millennial workforce’s reported penchant for job-hopping, Freshii boasts a headquarters team that is 98% millennial with extremely low turnover. A millennial herself, Ashley Dalziel attributes this success to the incredible appeal the company’s mission holds for her generation: to promote healthy living by providing affordable and convenient healthy food. “People want to do work that matters,” she says. “They want more than just a job and want to be part of a company that is about more than just making a profit.” Dalziel has a strong background in recruitment, employment branding, talent management and leadership development, having spent a decade at Lululemon Athletica, where she helped it grow into a multi-billion-dollar organization. Her work at Freshii is already paying dividends, too – the company won the Best Workplace Culture Award at the 2016 Canadian HR Awards.
ANNA PETOSA
CAVELL FRASER
Vice-president, people and culture
LIBRO CREDIT UNION
Senior HR consultant
PELMOREX MEDIA
As VP of people and culture for a media company approaching its 30-year mark, Anna Petosa’s top priority for the past year has been spearheading a cultural transformation project to ensure Pelmorex keeps pace with constant disruptions in the industry. In response to one identified HR service gap – the need for employees to share the vision of the company and see the relevance of their roles – Petosa and her team launched a program called ROAD Trip to a Performance Culture to increase engagement and foster the desired performance culture. The program is composed of four aspects: Recognition (of others and its importance), Objectives (setting SMART ones), Assessment (regular dialogue with managers) and Development (continuous L&D). For 2017, Petosa will be focusing on harnessing HR data analytics to completely shift Pelmorex Media’s HR department from an administrative role to a strategic one.
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Cavell Fraser has led Libro Credit Union’s HR team to support its new business strategy of ‘Growing Prosperity in Southwestern Ontario.’ This strategy incorporated two mergers, one in 2014 and another in 2015. Under Fraser’s leadership, the team has achieved a number of milestones, including completing two banking conversions, managing 154 internal job placements among 640 staff and hiring 123 new employees, including those from the merger. In 2016, Fraser completed two key initiatives: the rollout of an Aon Hewitt engagement survey, which led to the company being named Best Employer (Platinum Level) for both 2016 and 2017, and the setup of a Human Resources Management System [HRMS] committee. Fraser helped conduct a needs analysis for a new HRMS in the organization and created a comparison report of the final vendors. Her next step is to lead the HR team to implement the system and conduct the necessary training in early 2017.
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COLLEEN BAILEY MOFFITT CHRO ENERCARE
Colleen Bailey Moffitt joined Enercare as CHRO in April 2015. By May 2016, following multiple acquisitions, the company had grown to nearly 3,800 employees. In the midst of this rapid growth, Moffitt and her team have been focused on maintaining a high-performance culture and ensuring an aligned employee experience through various HR processes such as talent and performance management, reward & recognition, and total engagement. In 2017, Moffitt is expecting to implement a new HRIS and continue embedding newly established programs. Moffitt has more than 20 years of experience in HR across various industries such as energy, finance, retail and consulting, and has held executive HR roles at major companies such as TD Bank, PwC and Direct Energy.
ELAINE CRUISE SMITH Chief talent and culture leader WORLDLYNX WIRELESS
Under Elaine Cruise Smith’s leadership, communications retailer Worldlynx was named a Great Place to Work in 2016, as well as a Best Workplace for Women in Canada, with an all-time-high employee satisfaction rating of 84.6%. Smith’s focus for the past year has been on wellness, employee development and diversity. On the diversity front, she launched an employee resource group for women’s mentoring and networking, and introduced various diversity initiatives as part of the Worldlynx recruitment strategy to attract not only women, but also millennials, new graduates, people with disabilities and at-risk youth. Her acute understanding of the company’s culture – which is grounded in core values – has also been used to develop sales tools that the company’s sales team relies on to win clients, providing Worldlynx with a clear example of how HR can contribute to the bottom line.
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FEATURES
COVER STORY: HOT LIST 2017 HEATHER WYLLIE Vice-president of people NANDO’S CANADA
In her current role as VP of people for Nando’s, Heather Wyllie ensures that close ties and relationships exist between the people leaders of various offices around the world. “It’s no secret to our Canadian business or to Nando’s globally that it’s never been about the chicken; it’s about the people who make the chicken … [and] Heather brings people to the forefront of our Canadian business,” writes a colleague. In just under two years with Nando’s Canada, Wyllie has led the growth of the central team and is continuously working with senior leadership to map out long-term strategic growth plans, which will be essential to the company’s goal to almost double in size. Wyllie has built talent from the ground up for Nando’s Canada: Over the past 12 months, the company has launched a talent strategy and supporting tools for more than 900 ‘Nandocas’ across more than 40 locations in three provinces.
ILKA BENE
JEFF WALDMAN
Assistant vice-president of HR
Founder
ISLAND SAVINGS (A DIVISION OF FIRST WEST CREDIT UNION)
SOCIALHRCAMP
Following the integration of Island Savings into First West Credit Union in 2015, Ilka Bene was instrumental in the communication and implementation of all people services-related decisions for Island Savings. She has led a number of change projects, including harmonizing policies, job titles, and the compensation and benefits program at Island Savings, in addition to overseeing software system conversions and leading union relations/collective bargaining. Most important, Bene worked to keep the balance between developing a shared culture and retaining a local culture, ensuring that policies and practices remained aligned with the parent organization while the implementation and outcomes were localized. To ensure that employee engagement remained a priority, Bene chaired a committee called Islands’ Voice, which served as the testing ground for new ideas, projects and policies. As a result, Island Savings’ engagement score for 2016 was the highest within First West’s regions.
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What’s New in the J-Perk 5.01 serial key or number?
Screen Shot
System Requirements for J-Perk 5.01 serial key or number
- First, download the J-Perk 5.01 serial key or number
-
You can download its setup from given links:
J-Perk 5.01 serial key or number & for Laptop
J-Perk 5.01 serial key or number& Apps for Laptop & PC Free Download
J-Perk 5.01 serial key or number
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