The potential health hazards of microwaves have attracted much more attention. Our previous study found that 2856 MHz microwave radiation damaged synaptic plasticity and activated autophagy in neurons. However, the mechanisms underlying microwave-induced autophagy were still unclear. In the present study, we established neuronal damage models by exposing rat hippocampal neurons and rat adrenal pheochromocytoma (PC12) cell-derived neuron-like cells to 30 mW/cm2 microwaves, which resulted in miR-30a-5p ('miR-30a' for short) down-regulation and autophagy activation in vivo and in vitro. Bioinformatics analysis was conducted, and Beclin1, Prkaa2, Irs1, Pik3r2, Rras2, Ddit4, Gabarapl2 and autophagy-related gene 12 (Atg12) were identified as potential downstream genes of miR-30a involved in regulating autophagy. Based on our previous findings that microwave radiation could lead to abnormal energy metabolism in neurons, Prkaa2, encoding adenosine 5'-monophosphate-activated protein kinase (AMPK) α2 (AMPKα2, an important catalytic subunit of energy sensor AMPK), was selected for further analysis. Dual-luciferase reporter assay results showed that Prkaa2 was a downstream gene of miR-30a. Moreover, microwave radiation increased the expression of AMPKα2 and the phosphorylation of AMPKα (Thr172) both in vivo and in vitro. The transfection of PC12 cells with miR-30a mimics increased miR-30a levels, reduced AMPKα2 expression, suppressed AMPKα (Thr172) phosphorylation, and inhibited autophagy occurrence in neuron-like cells. Importantly, miR-30a overexpression abolished microwave-activated autophagy and inhibited microwave-induced AMPKα2 up-regulation and AMPKα (Thr172) phosphorylation. In conclusion, microwave radiation promoted the occurrence of autophagy in neurons through the miR-30a/AMPKα2 signal pathway.
The potential health hazards of microwaves have attracted much more attention. Our previous study found that 2856 MHz microwave radiation damaged synaptic plasticity and activated autophagy in neurons. However, the mechanisms underlying microwave-induced autophagy were still unclear. In the present study, we established neuronal damage models by exposing rat hippocampal neurons and rat adrenal pheochromocytoma (PC12) cell-derived neuron-like cells to 30 mW/cm2 microwaves, which resulted in miR-30a-5p ('miR-30a' for short) down-regulation and autophagy activation in vivo and in vitro. Bioinformatics analysis was conducted, and Beclin1, Prkaa2, Irs1, Pik3r2, Rras2, Ddit4, Gabarapl2 and autophagy-related gene 12 (Atg12) were identified as potential downstream genes of miR-30a involved in regulating autophagy. Based on our previous findings that microwave radiation could lead to abnormal energy metabolism in neurons, Prkaa2, encoding adenosine 5'-monophosphate-activated protein kinase (AMPK) α2 (AMPKα2, an important catalytic subunit of energy sensor AMPK), was selected for further analysis. Dual-luciferase reporter assay results showed that Prkaa2 was a downstream gene of miR-30a. Moreover, microwave radiation increased the expression of AMPKα2 and the phosphorylation of AMPKα (Thr172) both in vivo and in vitro. The transfection of PC12 cells with miR-30a mimics increased miR-30a levels, reduced AMPKα2 expression, suppressed AMPKα (Thr172) phosphorylation, and inhibited autophagy occurrence in neuron-like cells. Importantly, miR-30a overexpression abolished microwave-activated autophagy and inhibited microwave-induced AMPKα2 up-regulation and AMPKα (Thr172) phosphorylation. In conclusion, microwave radiation promoted the occurrence of autophagy in neurons through the miR-30a/AMPKα2 signal pathway.
Microwave use is widespread in modern life. Radiofrequency, including microwaves, had been classified as 2B carcinogens in 2013. Moreover, the National Toxicology Program of U.S.A. has reported the latest evidence that microwave radiation with a frequency of 900 MHz can cause cancers such as glioma in rats [1]. The brain is considered to be sensitive to microwave radiation, as many studies have reported the damaging effects of microwave exposure on the brain [2-5]. Therefore, it is important to study the biological effects and underlying mechanisms of microwave-induced neuronal damage. Autophagy, typically referred to as macroautophagy, is a cellular process that sequesters, removes and recycles unwanted macromolecules and damaged organelles, and is of great importance in maintaining cellular homeostasis [6]. Numerous studies have shown that autophagy plays important roles in the pathophysiological processes of various diseases, including tumors and neurodegenerative disorders [7-11]. In our previous study, autophagy was found to be activated in rat hippocampal neurons following microwave exposure [12]. However, the mechanisms involved in the regulation of autophagy induced by microwave radiation remain unclear.MicroRNAs (miRNAs) are 20–24 nucleotide noncoding RNAs that can affect protein expression at the post-transcriptional level by modulating the stability and translation of corresponding messenger RNAs (mRNAs) [13]. The roles played by miRNAs in various biological processes have received increasing attention. In our previous study, several differentially expressed miRNAs were screened from the rat hippocampus exposed to microwave radiation [14], among which miR-30a-5p (‘miR-30a’ for short) attracted our attention due to its regulatory effects on autophagy. Studies have reported that miR-30a negatively regulates Beclin1-mediated autophagy, which has emerged as a promising therapeutic target for multiple diseases, such as cerebral ischemic stroke, infection and cancers [15-19]. Additionally, we previously found that microwave radiation could induce abnormal energy metabolism in neurons [20,21]. As an energy sensor, adenosine 5′-monophosphate-activated protein kinase (AMPK) plays key roles in maintaining cellular metabolic balance and in regulating autophagy [22]. However, whether and how miR-30a and AMPK regulated microwave-mediated autophagy activation requires further elucidation.In the present study, we showed that microwave radiation promoted autophagy occurrence and reduced miR-30a expression both in rat hippocampal neurons and rat adrenal pheochromocytoma (PC12) cell-derived neuron-like cells. Prkaa2, encoding AMPKα2, was identified as one of the downstream genes of miR-30a. Microwave radiation increased AMPKα2 expression and AMPKα (Thr172) phosphorylation both in vivo and in vitro. In neuron-like cells, miR-30a overexpression abolished microwave-induced AMPKα2 up-regulation and autophagy activation.
Materials and methods
Animals and microwave exposure
All animal experiments took place at the Beijing Institute of Radiation Medicine. All experimental procedures were performed in accordance with the National Institutes of Health Guide for the Care and Use of Laboratory Animals and were approved by the Institutional Animal Care and Use Committee of the Beijing Institute of Radiation Medicine.Sample size was arbitrarily set to 48 (two groups with 24 animals each). Male Wistar rats (212.5 ± 7.1 g, 8-week-old, specific pathogen-free) were provided by and maintained in the Laboratory Animal Center of Beijing Institute of Radiation Medicine, where the temperature was 22 ± 2°C and the humidity 55 ± 10% on a 12-h light–dark cycle. Food and water were freely available. The rats were randomly divided into two groups: the microwave-exposed (MW) and sham-exposed (SH) groups. As described in our previous study [12], the rats in the MW group were placed in fan-shaped boxes made of plexiglass and free of metal just below the microwave source and exposed to 2856 MHz microwaves at an average power density of 30 mW/cm2 for 15 min, once every other day for three exposures. The specific absorption rate (SAR) was approximately 10.5 W/kg; the calculation method has been described previously [23]. The rats in the SH group were processed in parallel with those in the MW group, but with the microwave source switched off. At appointed time points, rats were anesthetized with 1% pentobarbital sodium (30 mg/kg) by intraperitoneal injection to minimize animal suffering during the procedure. The rats were then decapitatedand and hippocampi were isolated and used for further tests.
Cell culture and microwave exposure
PC12 cells, provided by China Infrastructure of Cell Line Resource, were cultured in RPMI 1640 basic medium (Gibco, Waltham, MA) supplemented with 10% horse serum (HS; Gibco) and 5% fetal bovine serum (FBS; Gibco). To induce the formation of neuron-like cells, PC12 cells were maintained in RPMI 1640 basic medium supplemented with 10 ng/ml nerve growth factor (NGF; Sigma, St. Louis, MO) and 1% HS for 5 days. Subsequently, the neuron-like cells with neuronal phenotype (an extension of neurites) were observed.The neuron-like cells were cultured in a six-well plate containing 2 ml medium, and randomly divided into the MW and SH groups. The cells in the MW group were exposed to 2856 MHz microwave radiation with an average power density of 30 mW/cm² for single 15 min. The SAR was calculated to be approximately 19.0 W/kg. Similar processing was conducted on cells in the SH group but without microwave radiation. Thereafter, the cells were harvested for analysis at specific time points.
Transmission electron microscopy
At 7 days, 14 days and 1 month, rat hippocampi were isolated and 1 mm3 tissue blocks were collected. At 6 h, neuron-like cells cultured on removable 96-well plate were harvested. Then, the samples were successively fixed in 2.5% glutaraldehyde and 1% osmium acid, processed with graded ethyl alcohols and embedded in EPON618. After being cut into ultrathin (70-nm) sections, the samples were subsequently stained with uranyl acetate and lead citrate. The ultrastructure of rat hippocampal neurons and neuron-like cells, especially autophagosomes and autolysosomes, was observed under transmission electron microscopy (TEM) (Hitachi, Japan).
Western blots
Total proteins were extracted from rat hippocampus at 7 days, 14 days and 1 month after exposure from neuron-like cells at 6, 12 and 24 h after exposure. Beclin1, microtubule-associated protein light chain 3 (MAP/LC3 or LC3), autophagy-related gene (Atg) 5 (Atg5), Atg7, Atg9, AMPKα2, p-AMPKα (Thr172), AMPKα and glyceraldehyde-3-phosphate dehydrogenase (GAPDH) were labeled with a rabbit anti-Beclin1 antibody (1:1000 dilution; #3,495; Cell Signaling Technology, CST), rabbit anti-LC3A/B antibody (1:1000 dilution; #12,741; CST), rabbit anti-Atg5 antibody (1:1000 dilution; #12,994; CST), rabbit anti-Atg7 antibody (1:1,00 dilution; #8,558; CST), rabbit anti-Atg9A antibody (1:1000 dilution; #13,509; CST), rabbit anti-AMPKα2 antibody (1:1000 dilution; #3,760; Abcam), rabbit anti-phospho-AMPKα (Thr172) antibody (1:1000 dilution; #2,535; CST), rabbit anti-AMPKα antibody (1:1000 dilution; #5,832; CST), and mouse anti-GAPDH antibody (1:5000 dilution; #8,245; Abcam), respectively. After incubating with the corresponding goat anti-rabbit IgG-horseradish peroxidase (HRP) (1:5000 dilution; #2,004; Santa Cruz) and goat anti-mouse IgG-HRP (1:5000 dilution; #2,005; Santa Cruz), the protein bands were recorded using an X-OMAT BT Film imaging system (Carestream, Rochester, NY). Image-Pro Plus 6.0 (Media Cybernetics, Rockville, MD) was used to analyze the integrated optical density (IOD) of protein bands.
Autophagic flux analysis after microwave exposure
To analyze autophagic flux in neuron-like cells after microwave exposure, 50 μM chloroquine (CQ, #142116, Abcam) was added 1 h before microwave radiation to inhibit the lysosome-mediated degradation of autophagosomes. LC3-II and LC3-I expression in cells pretreated with or without CQ was detected by Western blot at 6, 12 and 24 h after microwave exposure, using rabbit anti-LC3A/B antibody (1:1000 dilution; #12,741; CST), and the ratio of LC3-II to GAPDH was calculated and statistically analyzed.The expression and localization of LC3 protein in neuron-like cells treated with or without CQ was detected by immunofluorescence (IF) staining. Briefly, neuron-like cells grown on coverslips were harvested at 6 h after exposure and fixed with a mixture of methanol and acetone (1:1) for 10 min at room temperature. Then, the cells were labeled with rabbit anti-LC3A/B antibody (1:100 dilution; #12,741; CST) overnight at 4°C. After being washed, the cells were incubated with fluorescein isothiocyante (FITC)-conjugated goat anti-rabbit IgG secondary antibody (1:200 dilution; #0311; ZSGB-BIO; Beijing, China) for 1 h at room temperature. The cell nuclei were stained with 4′,6-diamidino-2-phenylindole (DAPI) for 5 min. Finally, the expression and localization of LC3 in cells was observed using a fluorescence microscope (DM6000, Leica, Wetzlar, Germany).
MiR-30a expression in rat hippocampi and neuron-like cells
At appointed time points after microwave radiation, rat hippocampi were collected and neuron-like cells were harvested. Total RNA was isolated from rat hippocampi and neuron-like cells using an mirVana miRNA Isolation kit (Ambion, Waltham, MA), according to the manufacturer’s instructions. Thereafter, complementary DNA (cDNA) was synthesized using a TaqMan MicroRNA Reverse Transcription kit (Applied Biosystems, ABI, Waltham, MA). The expression of miR-30a was quantified by real-time reverse-transcript polymerase chain reaction (RT-PCR) using TaqMan Universal Master Mix II (with UNG) and TaqMan MicroRNA assays (ABI). Finally, the expression of miR-30a was normalized to that of U6.
In situ hybridization
At 7 and 14 days after microwave radiation, rat brains were collected and fixed in 10% buffered formalin solution, embedded in paraffin and cut into 3-μm-thick sections. Then, miR-30a expression was assessed by in situ hybridization (ISH). Briefly, the sections were hybridized with miR-30a, U6, or scramble miRNA probes (Exiqon, Vedbaek, Denmark). After being labeled with anti-DIG (Boster Biological Technology, China) and stained with diaminobenzidine (DAB) (ZSGB-Bio), the expression of miR-30a and U6 was assessed in a blind manner under a light microscope (Leica). To quantify miR-30a expression, Image-Pro Plus 6.0 was used to analyze the IOD, and the expression of miR-30a was normalized to that of U6.
Bioinformatics analysis of downstream target genes of miR-30a
To predict the target genes of miR-30a, miRanda (Version 1.9, http://cbio.mskcc.org/miRNA2003/miranda.html), miRDB (Version 5.0, http://mirdb.org/index.html), TargetScan (Version 7.1, http://www.targetscan.org/mmu_71/), and miWalk (Version 2.0, http://zmf.umm.uni-heidelberg.de/apps/zmf/mirwalk2/) analysis was conducted, and only the genes co-predicted at least by three databases were selected as candidates for further analysis. Gene naming was standardized using NCBI Entrez Gene ID. The functional enrichment analysis of the target genes of miR-30a was conducted using clusterProfiler [24] with the Kyoto Encyclopedia of Genes and Genomes.db (KEGG.db; Version 83.1, http://www.genome.jp/kegg/) [25]. Significance was assessed using hypergeometric tests and Benjamini–Hochberg correction, with a significance threshold of P<0.05.
Luciferase reporter vectors and luciferase assay
HEK-293T cells were transiently transfected with pmirGLO-rPrkaa2 3′-untranslated region (3′UTR) wildtype (WT, 5′-CCUUCUGUUUACUUUUAGAA-3′), pmirGLO-rPrkaa2 3′UTR mutant type (MUT, 5′-CCUUCCAAUAGAUUUUAGAA-3′) and empty pmirGLO vectors (GenePharma, Shanghai, China) in combination with miR-30a mimics/negative control miRNAs (Ambion), using Lipofectamine 2000 (Invitrogen). Cells were lysed by Passive Lysis Buffer (Promega, Madison, WI) for 15 min. Then, firefly and Renilla luciferase activities were detected using a Dual Luciferase Assay kit (Promega) on a luminometer (Thermo Fisher Scientific).
RNA transfection
At 24 h before microwave exposure, neuron-like cells were transiently transfected with 12.5 nM miR-30a mimics and negative control miRNAs (Ambion) using Lipofectamine RNAiMAX (Invitrogen). Neuron-like cells were harvested at 6 h after microwave exposure. The expression of miR-30a was analyzed by real-time RT-PCR. The protein levels of AMPKα2, p-AMPKα (Thr172), AMPKα, Beclin1, LC3-I, LC3-II and GAPDH were detected by Western blots.
Statistical analysis
Data are presented as the mean ± standard error of the mean (SEM). All experiments were performed with a minimum of three independent replicates. The statistical analysis was achieved by the SPSS software (IBM, Armonk, NY, U.S.A.). For the data from two groups SH and MW with different time points, one-way analysis of variance (ANOVA) with repeated measures was used to analyze the time course, then Student’s t test was used to compare the differences between groups. For experiments of two-factor factorial design, two-way ANOVA was performed to analyze the effects of two factors. Besides, one-way ANOVA followed by Bonferroni’s post-hoc tests were performed to compare multiple groups. The differences were considered significant at the level of a two-sided P<0.05.
Results
Microwave radiation promoted the occurrence of autophagy in rat hippocampal neurons and PC12 cell-derived neuron-like cells
First, the ultrastructure of rat hippocampal neurons was observed by TEM at 7 days, 14 days and 1 month after 30 mW/cm2 microwave radiation. The autophagosomes can be identified by its contents (morphologically intact cytoplasm), and the limiting membrane that is partially visible as two bilayers separated by a narrow electron-lucent cleft. The autolysosomes can be identified by its contents, partially degraded, electron-dense rough endoplasmic reticulum [26]. The number of autophagosomes and autolysosomes in neurons notably increased from 7 days to 1 month after microwave exposure. Double-membrane autophagosomes, encapsulating mitochondria, synaptic vesicles and other cytosolic constituents, were primarily distributed at the synaptic terminal of hippocampal neurons. Monolayer autolysosomes, shown as black granular or amorphous aggregates, degraded cytoplasmic components at different stages and were mainly located in cell bodies of hippocampal neurons (Figure 1A–C). To analyze the dynamic regulation of microwave radiation on autophagy, Beclin1 and LC3, two widely used molecular markers of autophagy, were analyzed in rat hippocampi. The expression of Beclin1 was up-regulated 14 days after microwave radiation, suggesting the activation of autophagy, but down-regulated at 1 month which might be attributed to a regulatory adjustment (Figure 1D,E). The expression of LC3-II significantly increased 14 days and 1 month after microwave exposure, indicating an increased number of autophagosomes (Figure 1F,G). These results suggested that 30 mW/cm2 microwave radiation promoted the occurrence of autophagy in rat hippocampal neurons.
Figure 1
Microwaves promoted autophagy occurrence in rat hippocampal neurons
(A) Representative images showing autophagosomes and autolysosomes in hippocampal neurons observed by TEM (scale bar = 500 nm), 7 days, 14 days and 1 month postradiation. The autophagosomes are indicated by arrows without tails, while the autolysosomes are indicated by arrows with tails. (B,C) The quantification results of the total number of autophagosomes and autolysosomes in synapse and cell body and statistical analysis. (D,E) Representative protein bands of Beclin1 and GAPDH in rat hippocampus and quantitative analysis result, on 7 days, 14 days and 1 month postradiation. (F,G) Representative protein bands of LC3-I, LC3-II and GAPDH in rat hippocampus and quantitative analysis result, on 7 days, 14 days and 1 month postradiation. The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups SH and MW (B,C,E,G). *, P<0.05; **, P<0.01 vs the control.
Microwaves promoted autophagy occurrence in rat hippocampal neurons
(A) Representative images showing autophagosomes and autolysosomes in hippocampal neurons observed by TEM (scale bar = 500 nm), 7 days, 14 days and 1 month postradiation. The autophagosomes are indicated by arrows without tails, while the autolysosomes are indicated by arrows with tails. (B,C) The quantification results of the total number of autophagosomes and autolysosomes in synapse and cell body and statistical analysis. (D,E) Representative protein bands of Beclin1 and GAPDH in rat hippocampus and quantitative analysis result, on 7 days, 14 days and 1 month postradiation. (F,G) Representative protein bands of LC3-I, LC3-II and GAPDH in rat hippocampus and quantitative analysis result, on 7 days, 14 days and 1 month postradiation. The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups SH and MW (B,C,E,G). *, P<0.05; **, P<0.01 vs the control.The changes of autophagy in PC12 cell-derived neuron-like cells after microwave exposure was also studied. To evaluate ‘autophagic flux’, the LC3-II levels was detected in neuron-like cells pretreated with or without CQ, a lysosomal inhibitor. In the CQ-untreated cells, the LC3-II content showed no difference between the SH and MW groups. When the degradation of autophagosomes by lysosomes was inhibited by CQ, the expression of LC3-II in cells from the MW group obviously increased at 6 h but restored to basal levels at 12 and 24 h after exposure, indicating the enhanced ‘autophagic flux’ in neuron-like cells induced by microwave radiation (Figure 2A,B). The expression of autophagy markers Beclin1 was up-regulated at 6 h and restored at 12 h postradiation, and Atg5, Atg7 and Atg9 were up-regulated at 6 and 12 h after microwave exposure, indicating the enhanced expression of autophagy-related genes induced by microwaves (Figure 2A,B). Furthermore, we observed that autophagosomes and autolysosomes in neuron-like cells showed an increasing trend at 6 h after microwave exposure (Figure 2C). Through IF staining, an increase in the number of LC3 puncta (autophagosomes) was observed in neuron-like cells pretreated with CQ at 6 h after microwave radiation (Figure 2D,E). Our data suggested that 30 mW/cm2 microwave radiation activated functional autophagy in neuron-like cells.
Figure 2
Microwave radiation activated functional autophagy in PC12 cell-derived neuron-like cells
(A) Representative protein bands of LC3-1, LC3-II, Beclin1, Atg5, Atg7, Atg9 and GAPDH in PC12 cell-derived neuron-like cells induced by NGF, at 6, 12 and 24 h after 30 mW/cm2 microwave exposure. To assess ‘autophagic flux’, neuron-like cells were pretreated with chloroquine (CQ+) or equal amount of water (CQ−) 1 h before microwave exposure, and the expression of LC3-1 and LC3-II was detected by Western blots. (B) The statistical analysis results of protein bands in (A). (C) Representative images showing autophagosomes and autolysosomes in neuron-like cells at 6 h after exposure, observed by TEM (scale bar = 500 nm). (D) Representative images of LC3 dots in neuron-like cells at 6 h after radiation, detected by IF (scale bar = 20 μm). ‘CQ pretreated’ indicated that the cells were treated with CQ 1 h before microwave radiation, while ‘CQ untreated’ meant the cells were treated with equal amount of water. (E) The statistical analysis results of LC3 dots in (D). The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups SH and MW (B). Two-way ANOVA was performed to analyze the data of two-factor factorial design (E). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control.
Microwave radiation activated functional autophagy in PC12 cell-derived neuron-like cells
(A) Representative protein bands of LC3-1, LC3-II, Beclin1, Atg5, Atg7, Atg9 and GAPDH in PC12 cell-derived neuron-like cells induced by NGF, at 6, 12 and 24 h after 30 mW/cm2 microwave exposure. To assess ‘autophagic flux’, neuron-like cells were pretreated with chloroquine (CQ+) or equal amount of water (CQ−) 1 h before microwave exposure, and the expression of LC3-1 and LC3-II was detected by Western blots. (B) The statistical analysis results of protein bands in (A). (C) Representative images showing autophagosomes and autolysosomes in neuron-like cells at 6 h after exposure, observed by TEM (scale bar = 500 nm). (D) Representative images of LC3 dots in neuron-like cells at 6 h after radiation, detected by IF (scale bar = 20 μm). ‘CQ pretreated’ indicated that the cells were treated with CQ 1 h before microwave radiation, while ‘CQ untreated’ meant the cells were treated with equal amount of water. (E) The statistical analysis results of LC3 dots in (D). The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups SH and MW (B). Two-way ANOVA was performed to analyze the data of two-factor factorial design (E). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control.
Microwave radiation inhibited miR-30a expression in rat hippocampi and neuron-like cells
It has been widely reported that miRNAs play important roles in physiological and pathological events in the central nervous system [27-29]. We previously found that microwave radiation could induce cognitive dysfunction and neuron injury in rats [12]. In another study, we screened the differentially expressed miRNAs in the rat hippocampus after microwave radiation, among which miR-30a attracted our interests due to its potential regulatory roles on autophagy (Figure 3A) [14]. Based on these results, in the present study, we confirmed the changes in miR-30a levels induced by microwave radiation both in vivo and in vitro. The results suggested that microwave radiation inhibited the expression of miR-30a both in rat hippocampi and neuron-like cells (Figure 3B,C). The decreased expression of miR-30a in rat hippocampi was subsequently confirmed by ISH (Figure 3D,E). These results indicated that microwave radiation inhibited miR-30a both in vivo and in vitro, which might be the important mechanisms underlying microwave-induced autophagy in neurons.
Figure 3
Microwave radiation inhibited miR-30a expression both in rat hippocampi and neuron-like cells
(A) Differentially expressed miRNAs in the rat hippocampi screened by miRNA chip. (B,C) The expression of miR-30a and U6 in rat hippocampi 7 days, 14 days and 1 month after radiation and PC12 cell-derived neuron-like cells 6, 12 and 24 h after radiation, respectively, which were detected by real-time RT-PCR. (D) Representative images of miR-30a and U6 in rat hippocampi 7 and 14 days after radiation, detected by ISH (scale bar = 30 μm). (E) The quantitative analysis result of the ratio of miR-30a to U6 presented in (D). The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups: SH and MW (B,C,E). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control.
Microwave radiation inhibited miR-30a expression both in rat hippocampi and neuron-like cells
(A) Differentially expressed miRNAs in the rat hippocampi screened by miRNA chip. (B,C) The expression of miR-30a and U6 in rat hippocampi 7 days, 14 days and 1 month after radiation and PC12 cell-derived neuron-like cells 6, 12 and 24 h after radiation, respectively, which were detected by real-time RT-PCR. (D) Representative images of miR-30a and U6 in rat hippocampi 7 and 14 days after radiation, detected by ISH (scale bar = 30 μm). (E) The quantitative analysis result of the ratio of miR-30a to U6 presented in (D). The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups: SH and MW (B,C,E). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control.
AMPKα2 was one of the downstream genes of miR-30a, microwave radiation increased AMPKα2 expression and AMPK (Thr172) phosphorylation
miR-30a has been shown to be an important molecule in the regulation of autophagy [15-18,30]. In the present study, we predicted the downstream genes of miR-30a using four databases, including miRanda, miRDB, TargetScan and miWalk. Subsequently, 365 target genes of miR-30a co-predicted by at least three databases were selected for KEGG analysis (Figure 4A). Our data showed that the predicted target genes of miR-30a were involved in multiple signal pathways, such as autophagy, cellular senescence, cytokine–cytokine receptor interactions, the cyclic guanosine monophosphate-dependent protein kinase (cGMP-PKG) signal pathway, and the apelin signal pathway (Figure 4B). As the potential downstream genes of miR-30a, Beclin1, Prkaa2, Irs1, Pik3r2, Rras2, Ddit4, Gabarapl2 and Atg12 appeared to play important roles in regulating autophagy. Studies have reported that Beclin1, Gabarapl2 and Atg12 mediate the induction and formation of autophagosomes. Prkaa2, encoding AMPKα2, may be involved in the regulation of autophagy through the AMPK signal pathway. REDD1, Irs1, Pik3r2, and Rras2 appears to regulate autophagy through the mammalian target of rapamycin (mTOR) signal pathway (Figure 4C) [31,32]. Among that, the role of Beclin1 in miR-30a-regulated autophagy has been widely studied [33-36]. In our research and from other scholars, microwave radiation can induce abnormal energy metabolism in neurons [20,21,37,38]. Studies have confirmed that microwave under certain conditions can promote the occurrence of autophagy, but the underlying mechanisms have yet to be clarified [12,39-42]. Accordingly, in the present study, we further explored the roles of Prkaa2 played in the miR-30a-mediated regulation of autophagy in MW neuron models, which is yet to be reported.
Figure 4
The downstream genes of miR-30a were predicted and analyzed through bioinformatics
(A) The numbers of the predicted downstream genes of miR-30a by four databases, including miRanda, miRDB, TargetScan and miWalk. (B) The signal pathways regulated by miR-30a analyzed by the KEGG database. (C) The potential downstream genes of miR-30a involved in the regulation of autophagy, including Beclin1, Prkaa2, Irs1, Pik3r2, Rras2, Ddit4, Gabarapl2, and Atg12 are labeled in red color.
The downstream genes of miR-30a were predicted and analyzed through bioinformatics
(A) The numbers of the predicted downstream genes of miR-30a by four databases, including miRanda, miRDB, TargetScan and miWalk. (B) The signal pathways regulated by miR-30a analyzed by the KEGG database. (C) The potential downstream genes of miR-30a involved in the regulation of autophagy, including Beclin1, Prkaa2, Irs1, Pik3r2, Rras2, Ddit4, Gabarapl2, and Atg12 are labeled in red color.The expression of AMPKα2 was up-regulated in rat hippocampi from 14 days to 1 month after microwave exposure (Figure 5A–C), as well as in neuron-like cells at 12 h after microwave exposure (Figure 5D,F). AMPKα phosphorylation at Thr172 is essential for the activation of AMPK, the level of which can better reflect the changes in the activity of AMPK signal pathway [43]. In rat hippocampi, p-AMPKα (Thr172) levels increased 14 days after microwave exposure (Figure 5A–C). In neuron-like cells, microwave radiation induced the up-regulation of p-AMPKα (Thr172) levels at 6 and 12 h after exposure (Figure 5D–F). Considering the previously observed changes in autophagy, AMPK signal appeared to exhibit the same pattern as that observed for autophagy after microwave radiation, both in vivo and in vitro. These results suggested that AMPKα2 encoded by Prkaa2 was involved in the regulation of autophagy in neurons after microwave radiation. In addition, we observed that the overexpression of miR-30a reduced the luciferase activity of Prkaa2 in 3′UTR WT reporter-transfected cells but not in Prkaa2 3′UTR MUT reporter-transfected cells, suggesting an upstream regulatory role of miR-30a on AMPKα2 (Figure 5G,H). Accordingly, we hypothesized that the effects of microwave radiation on neuronal autophagy might be achieved through the miR-30a/Prkaa2/AMPKα2 pathway.
Figure 5
Microwave radiation increased AMPKα2 expression and AMPKα (Thr172) phosphorylation in rat and PC12 cell models
(A) Representative protein bands of AMPKα2, p-AMPKα (Thr172), AMPKα and GAPDH in rat hippocampi on 7 days, 14 days and 1 month postradiation. (B,C) The statistical analysis results of the ratio of AMPKα2 to GAPDH and p-AMPKα (Thr172) to AMPKα shown in (A), respectively. (D) Representative protein bands of AMPKα2, p-AMPKα (Thr172), AMPKα and GAPDH in neuron-like cells at 6, 12 and 24 h postradiation. (E,F) The statistical analysis results of the ratio of AMPKα2 to GAPDH and p-AMPKα (Thr172) to AMPKα shown in (D), respectively. (G) The base sequences of pmirGLO-rPrkaa2 3′UTR WT, miR-30a, and pmirGLO-rPrkaa2 3′UTR MUT. (H) The ratio of firefly to Renilla luciferase activity in HEK-293T cells transiently transfected with pmirGLO-rPrkaa2 3′UTR WT, pmirGLO-rPrkaa2 3′UTR MUT and empty pmirGLO vectors in combination with miR-30a mimics or corresponding negative control miRNAs. The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups SH and MW (B,C,E,F). One-way ANOVA followed by Bonferroni’s post-hoc tests was used to compare multiple groups in (H). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control.
Microwave radiation increased AMPKα2 expression and AMPKα (Thr172) phosphorylation in rat and PC12 cell models
(A) Representative protein bands of AMPKα2, p-AMPKα (Thr172), AMPKα and GAPDH in rat hippocampi on 7 days, 14 days and 1 month postradiation. (B,C) The statistical analysis results of the ratio of AMPKα2 to GAPDH and p-AMPKα (Thr172) to AMPKα shown in (A), respectively. (D) Representative protein bands of AMPKα2, p-AMPKα (Thr172), AMPKα and GAPDH in neuron-like cells at 6, 12 and 24 h postradiation. (E,F) The statistical analysis results of the ratio of AMPKα2 to GAPDH and p-AMPKα (Thr172) to AMPKα shown in (D), respectively. (G) The base sequences of pmirGLO-rPrkaa2 3′UTR WT, miR-30a, and pmirGLO-rPrkaa2 3′UTR MUT. (H) The ratio of firefly to Renilla luciferase activity in HEK-293T cells transiently transfected with pmirGLO-rPrkaa2 3′UTR WT, pmirGLO-rPrkaa2 3′UTR MUT and empty pmirGLO vectors in combination with miR-30a mimics or corresponding negative control miRNAs. The data are presented as mean ± SEM. One-way ANOVA with repeated measures was used to analyze the time course, and Student’s t test was used to compare the difference between the two groups SH and MW (B,C,E,F). One-way ANOVA followed by Bonferroni’s post-hoc tests was used to compare multiple groups in (H). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control.
MiR-30a overexpression inhibited AMPKα2 up-regulation, AMPKα (Thr172) phosphorylation, and autophagy activation mediated by microwave radiation in cell model
To investigate the underlying mechanisms that microwave-induced autophagy enhancement in neurons, we transfected miR-30a mimics into neuron-like cells 24 h before microwave exposure. It was found that miR-30a mimics significantly increased the expression level of miR-30a in neuron-like cells, suggesting that the intervention measures were effective (Figure 6A).
Figure 6
Transfection of miR-30a mimics increased miR-30a levels, reduced AMPKα2 expression, suppressed AMPKα (Thr172) phosphorylation and inhibited autophagy in neuron-like cells induced by microwaves
(A) The expression of miR-30a in PC12 cell-derived neuron-like cells measured by RT-PCR. (B–F) Representative protein bands of AMPKα2, p-AMPKα (Thr172), AMPKα, Beclin1, LC3-I, LC3-II and corresponding GAPDHs, and related statistical analysis results. The cells were transfected with miR-30a mimics or related negative control miRNAs 24 h before 30 mW/cm2 microwave radiation, and harvested 6 h after exposure. CQ+ indicated that the neuron-like cells were pretreated with CQ 1 h before microwave exposure, while CQ− indicated that equal amount of water was added. The data are presented as mean ± SEM. Two-way ANOVA was performed to analyze the data of two-factor factorial design (A–F). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control. The data in panels (D,E) are from the same sample, so they share the same GAPDH.
Transfection of miR-30a mimics increased miR-30a levels, reduced AMPKα2 expression, suppressed AMPKα (Thr172) phosphorylation and inhibited autophagy in neuron-like cells induced by microwaves
(A) The expression of miR-30a in PC12 cell-derived neuron-like cells measured by RT-PCR. (B–F) Representative protein bands of AMPKα2, p-AMPKα (Thr172), AMPKα, Beclin1, LC3-I, LC3-II and corresponding GAPDHs, and related statistical analysis results. The cells were transfected with miR-30a mimics or related negative control miRNAs 24 h before 30 mW/cm2 microwave radiation, and harvested 6 h after exposure. CQ+ indicated that the neuron-like cells were pretreated with CQ 1 h before microwave exposure, while CQ− indicated that equal amount of water was added. The data are presented as mean ± SEM. Two-way ANOVA was performed to analyze the data of two-factor factorial design (A–F). *, P<0.05; **, P<0.01; ***, P<0.001 vs the control. The data in panels (D,E) are from the same sample, so they share the same GAPDH.Few studies have reported the regulatory roles of miR-30a on the AMPK signal. In our study, miR-30a overexpression mediated by mimic transfection decreased AMPKα2 expression and inhibited AMPKα (Thr172) phosphorylation in neuron-like cells, indicating that miR-30a negatively regulated the activity of AMPK signal. Meanwhile, the overexpressed miR-30a inhibited microwave-induced AMPK activation (Figure 6B,C). Therefore, microwave radiation could modulate the activity of AMPK signal by suppressing miR-30a.Furthermore, we found that miR-30a overexpression reduced Beclin1 levels in neuron-like cells and inhibited microwave-induced up-regulation of Beclin1 expression (Figure 6D). Importantly, the overexpressed miR-30a suppressed the ‘autophagic flux’ in neuron-like cells, and completely abolished microwave-induced enhancement of autophagy (Figure 6E,F). Taken together, microwave-induced autophagy in neurons was abrogated by miR-30a overexpression, and the effects of microwave on autophagy were achieved by the miR-30a/AMPK pathway.
Discussion
Autophagy has been reported to play dual roles in pathological processes of neurodegenerative diseases. At the early stage, autophagy can enhance the degradation of denatured proteins to maintain neuronal functions, whereas at the late stage, continuous activation of autophagy ultimately induces the autophagic cell death of neurons [44-48]. Many studies have demonstrated that autophagy participates in the pathological processes of electromagnetic radiation-induced damaging effects [12,39-42]. Additionally, we previously found that autophagy-mediated degradation of synaptic vesicles was a potential mechanism of synaptic plasticity injury caused by microwaves. However, the underlying mechanisms how microwaves affect autophagy activity are yet to be clarified. In the present study, we exposed rats and PC12 cell-derived neuron-like cells to 2856 MHz microwaves with an average power density of 30 mW/cm2. The SAR value of the microwave radiation on rats was calculated to be approximately 10.5 W/kg, and the SAR value of neuron-like cells was 19.0 W/kg. We found that microwave radiation in the present study promoted autophagy in both rat hippocampal neurons and neuron-like cells. It seemed that autophagy occurred earlier in neuron-like cells with a larger SAR than in rat hippocampal neurons. The results of our previous study showed that hippocampal neurons had the ability to initiate self-recovery after 30 mW/cm2 microwave exposure [21]. Accordingly, we speculated that neuronal autophagy was most likely a protective response to microwave radiation in this study. At higher exposure dose levels, microwave radiation might overactivate autophagy, resulting in an unbalanced cellular homeostasis and ultimately leading to the irreversible injury of neurons, although further study was noted to be required [49-51].Numerous studies have demonstrated that miR-30a negatively regulates autophagy, a process that is closely associated with the pathophysiological processes of multiple diseases, such as cerebral ischemic stroke, cancer and hepatic fibrosis [33-36]. However, the roles played by miR-30a in microwave-induced neuron injury and autophagy activation are yet to be explored. In the present study, we observed that 30 mW/cm2 microwave radiation inhibited the expression of miR-30a both in vivo and in vitro. Based on these results, we speculated that miR-30a might participate in the regulation of microwave-induced autophagy in neurons. The results of the present study showed that miR-30a levels were significantly down-regulated in rat hippocampal neurons from 7 days to 1 month after microwave radiation, at which time autophagy was notably activated. In addition, similar results were obtained for neuron-like cells. Furthermore, miR-30a overexpression completely abolished microwave-induced autophagy in neuron-like cells. Taken together, these results suggested that reduced miR-30 levels were an important underlying mechanism involved in the activation of autophagy mediated by microwaves in neurons.As an important molecular target that regulates autophagy, the role of miR-30a in Beclin1-mediated autophagy has been widely studied and reported for a variety of diseases [33-36]. The results of the present study also support that decreased miR-30a levels positively regulates autophagy by promoting Beclin1 expression in neurons exposed to microwaves. However, the potential downstream target genes of miR-30a involved in regulating autophagy have yet to be systematically studied. Through bioinformatics analysis, we predicted the target genes of miR-30a that might be involved in autophagy regulation, including Beclin1, Prkaa2, Irs1, Pik3r2, Rras2, Ddit4, Gabarapl2 and Atg12. Studies have demonstrated that microwave radiation leads to abnormal energy metabolism in neurons [20,21,37,38]. In the present study, we showed that microwave radiation activated AMPK signal, a cellular energy sensor, both in rat hippocampi and neuron-like cells. There is strong consensus that AMPK signal mediates the activation of autophagy in various cells [52]. We confirmed that Prkaa2, encoding AMPKα2, was a downstream gene of miR-30a, which was verified by double luciferase reporter assay results. Moreover, microwave radiation increased AMPKα2 expression and phosphorylation (Thr172) both in vivo and in vitro. And miR-30a overexpression inhibited the microwave-induced activation of AMPKα2 and p-AMPKα, and abolished microwave-induced autophagy in neuron-like cells. Therefore, with the exception of Beclin1, we found that the reduced miR-30a levels after microwave exposure promoted the occurrence of autophagy by activating the AMPK signal, which had not been previously reported. Overall, in MW animal and cell model used in the study, the downregulation of miR-30a induced by microwaves increased the expression of AMPKα2 and the activation of AMPK signaling, which promoted the occurrence of autophagy and the biogenesis of autophagosomes.
Authors: Daniel J Klionsky; Kotb Abdelmohsen; Akihisa Abe; Md Joynal Abedin; Hagai Abeliovich; Abraham Acevedo Arozena; Hiroaki Adachi; Christopher M Adams; Peter D Adams; Khosrow Adeli; Peter J Adhihetty; Sharon G Adler; Galila Agam; Rajesh Agarwal; Manish K Aghi; Maria Agnello; Patrizia Agostinis; Patricia V Aguilar; Julio Aguirre-Ghiso; Edoardo M Airoldi; Slimane Ait-Si-Ali; Takahiko Akematsu; Emmanuel T Akporiaye; Mohamed Al-Rubeai; Guillermo M Albaiceta; Chris Albanese; Diego Albani; Matthew L Albert; Jesus Aldudo; Hana Algül; Mehrdad Alirezaei; Iraide Alloza; Alexandru Almasan; Maylin Almonte-Beceril; Emad S Alnemri; Covadonga Alonso; Nihal Altan-Bonnet; Dario C Altieri; Silvia Alvarez; Lydia Alvarez-Erviti; Sandro Alves; Giuseppina Amadoro; Atsuo Amano; Consuelo Amantini; Santiago Ambrosio; Ivano Amelio; Amal O Amer; Mohamed Amessou; Angelika Amon; Zhenyi An; Frank A Anania; Stig U Andersen; Usha P Andley; Catherine K Andreadi; Nathalie Andrieu-Abadie; Alberto Anel; David K Ann; Shailendra Anoopkumar-Dukie; Manuela Antonioli; Hiroshi Aoki; Nadezda Apostolova; Saveria Aquila; Katia Aquilano; Koichi Araki; Eli Arama; Agustin Aranda; Jun Araya; Alexandre Arcaro; Esperanza Arias; Hirokazu Arimoto; Aileen R Ariosa; Jane L Armstrong; Thierry Arnould; Ivica Arsov; Katsuhiko Asanuma; Valerie Askanas; Eric Asselin; Ryuichiro Atarashi; Sally S Atherton; Julie D Atkin; Laura D Attardi; Patrick Auberger; Georg Auburger; Laure Aurelian; Riccardo Autelli; Laura Avagliano; Maria Laura Avantaggiati; Limor Avrahami; Suresh Awale; Neelam Azad; Tiziana Bachetti; Jonathan M Backer; Dong-Hun Bae; Jae-Sung Bae; Ok-Nam Bae; Soo Han Bae; Eric H Baehrecke; Seung-Hoon Baek; Stephen Baghdiguian; Agnieszka Bagniewska-Zadworna; Hua Bai; Jie Bai; Xue-Yuan Bai; Yannick Bailly; Kithiganahalli Narayanaswamy Balaji; Walter Balduini; Andrea Ballabio; Rena Balzan; Rajkumar Banerjee; Gábor Bánhegyi; Haijun Bao; Benoit Barbeau; Maria D Barrachina; Esther Barreiro; Bonnie Bartel; Alberto Bartolomé; Diane C Bassham; Maria Teresa Bassi; Robert C Bast; Alakananda Basu; Maria Teresa Batista; Henri Batoko; Maurizio Battino; Kyle Bauckman; Bradley L Baumgarner; K Ulrich Bayer; Rupert Beale; Jean-François Beaulieu; George R Beck; Christoph Becker; J David Beckham; Pierre-André Bédard; Patrick J Bednarski; Thomas J Begley; Christian Behl; Christian Behrends; Georg Mn Behrens; Kevin E Behrns; Eloy Bejarano; Amine Belaid; Francesca Belleudi; Giovanni Bénard; Guy Berchem; Daniele Bergamaschi; Matteo Bergami; Ben Berkhout; Laura Berliocchi; Amélie Bernard; Monique Bernard; Francesca Bernassola; Anne Bertolotti; Amanda S Bess; Sébastien Besteiro; Saverio Bettuzzi; Savita Bhalla; Shalmoli Bhattacharyya; Sujit K Bhutia; Caroline Biagosch; Michele Wolfe Bianchi; Martine Biard-Piechaczyk; Viktor Billes; Claudia Bincoletto; Baris Bingol; Sara W Bird; Marc Bitoun; Ivana Bjedov; Craig Blackstone; Lionel Blanc; Guillermo A Blanco; Heidi Kiil Blomhoff; Emilio Boada-Romero; Stefan Böckler; Marianne Boes; Kathleen Boesze-Battaglia; Lawrence H Boise; Alessandra Bolino; Andrea Boman; Paolo Bonaldo; Matteo Bordi; Jürgen Bosch; Luis M Botana; Joelle Botti; German Bou; Marina Bouché; Marion Bouchecareilh; Marie-Josée Boucher; Michael E Boulton; Sebastien G Bouret; Patricia Boya; Michaël Boyer-Guittaut; Peter V Bozhkov; Nathan Brady; Vania Mm Braga; Claudio Brancolini; Gerhard H Braus; José M Bravo-San Pedro; Lisa A Brennan; Emery H Bresnick; Patrick Brest; Dave Bridges; Marie-Agnès Bringer; Marisa Brini; Glauber C Brito; Bertha Brodin; Paul S Brookes; Eric J Brown; Karen Brown; Hal E Broxmeyer; Alain Bruhat; Patricia Chakur Brum; John H Brumell; Nicola Brunetti-Pierri; Robert J Bryson-Richardson; Shilpa Buch; Alastair M Buchan; Hikmet Budak; Dmitry V Bulavin; Scott J Bultman; Geert Bultynck; Vladimir Bumbasirevic; Yan Burelle; Robert E Burke; Margit Burmeister; Peter Bütikofer; Laura Caberlotto; Ken Cadwell; Monika Cahova; Dongsheng Cai; Jingjing Cai; Qian Cai; Sara Calatayud; Nadine Camougrand; Michelangelo Campanella; Grant R Campbell; Matthew Campbell; Silvia Campello; Robin Candau; Isabella Caniggia; Lavinia Cantoni; Lizhi Cao; Allan B Caplan; Michele Caraglia; Claudio Cardinali; Sandra Morais Cardoso; Jennifer S Carew; Laura A Carleton; Cathleen R Carlin; Silvia Carloni; Sven R Carlsson; Didac Carmona-Gutierrez; Leticia Am Carneiro; Oliana Carnevali; Serena Carra; Alice Carrier; Bernadette Carroll; Caty Casas; Josefina Casas; Giuliana Cassinelli; Perrine Castets; Susana Castro-Obregon; Gabriella Cavallini; Isabella Ceccherini; Francesco Cecconi; Arthur I Cederbaum; Valentín Ceña; Simone Cenci; Claudia Cerella; Davide Cervia; Silvia Cetrullo; Hassan Chaachouay; Han-Jung Chae; Andrei S Chagin; Chee-Yin Chai; Gopal Chakrabarti; Georgios Chamilos; Edmond Yw Chan; Matthew Tv Chan; Dhyan Chandra; Pallavi Chandra; Chih-Peng Chang; Raymond Chuen-Chung Chang; Ta Yuan Chang; John C Chatham; Saurabh Chatterjee; Santosh Chauhan; Yongsheng Che; Michael E Cheetham; Rajkumar Cheluvappa; Chun-Jung Chen; Gang Chen; Guang-Chao Chen; Guoqiang Chen; Hongzhuan Chen; Jeff W Chen; Jian-Kang Chen; Min Chen; Mingzhou Chen; Peiwen Chen; Qi Chen; Quan Chen; Shang-Der Chen; Si Chen; Steve S-L Chen; Wei Chen; Wei-Jung Chen; Wen Qiang Chen; Wenli Chen; Xiangmei Chen; Yau-Hung Chen; Ye-Guang Chen; Yin Chen; Yingyu Chen; Yongshun Chen; Yu-Jen Chen; Yue-Qin Chen; Yujie Chen; Zhen Chen; Zhong Chen; Alan Cheng; Christopher Hk Cheng; Hua Cheng; Heesun Cheong; Sara Cherry; Jason Chesney; Chun Hei Antonio Cheung; Eric Chevet; Hsiang Cheng Chi; Sung-Gil Chi; Fulvio Chiacchiera; Hui-Ling Chiang; Roberto Chiarelli; Mario Chiariello; Marcello Chieppa; Lih-Shen Chin; Mario Chiong; Gigi Nc Chiu; Dong-Hyung Cho; Ssang-Goo Cho; William C Cho; Yong-Yeon Cho; Young-Seok Cho; Augustine Mk Choi; Eui-Ju Choi; Eun-Kyoung Choi; Jayoung Choi; Mary E Choi; Seung-Il Choi; Tsui-Fen Chou; Salem Chouaib; Divaker Choubey; Vinay Choubey; Kuan-Chih Chow; Kamal Chowdhury; Charleen T Chu; Tsung-Hsien Chuang; Taehoon Chun; Hyewon Chung; Taijoon Chung; Yuen-Li Chung; Yong-Joon Chwae; Valentina Cianfanelli; Roberto Ciarcia; Iwona A Ciechomska; Maria Rosa Ciriolo; Mara Cirone; Sofie Claerhout; Michael J Clague; Joan Clària; Peter Gh Clarke; Robert Clarke; Emilio Clementi; Cédric Cleyrat; Miriam Cnop; Eliana M Coccia; Tiziana Cocco; Patrice Codogno; Jörn Coers; Ezra Ew Cohen; David Colecchia; Luisa Coletto; Núria S Coll; Emma Colucci-Guyon; Sergio Comincini; Maria Condello; Katherine L Cook; Graham H Coombs; Cynthia D Cooper; J Mark Cooper; Isabelle Coppens; Maria Tiziana Corasaniti; Marco Corazzari; Ramon Corbalan; Elisabeth Corcelle-Termeau; Mario D Cordero; Cristina Corral-Ramos; Olga Corti; Andrea Cossarizza; Paola Costelli; Safia Costes; Susan L Cotman; Ana Coto-Montes; Sandra Cottet; Eduardo Couve; Lori R Covey; L Ashley Cowart; Jeffery S Cox; Fraser P Coxon; Carolyn B Coyne; Mark S Cragg; Rolf J Craven; Tiziana Crepaldi; Jose L Crespo; Alfredo Criollo; Valeria Crippa; Maria Teresa Cruz; Ana Maria Cuervo; Jose M Cuezva; Taixing Cui; Pedro R Cutillas; Mark J Czaja; Maria F Czyzyk-Krzeska; Ruben K Dagda; Uta Dahmen; Chunsun Dai; Wenjie Dai; Yun Dai; Kevin N Dalby; Luisa Dalla Valle; Guillaume Dalmasso; Marcello D'Amelio; Markus Damme; Arlette Darfeuille-Michaud; Catherine Dargemont; Victor M Darley-Usmar; Srinivasan Dasarathy; Biplab Dasgupta; Srikanta Dash; Crispin R Dass; Hazel Marie Davey; Lester M Davids; David Dávila; Roger J Davis; Ted M Dawson; Valina L Dawson; Paula Daza; Jackie de Belleroche; Paul de Figueiredo; Regina Celia Bressan Queiroz de Figueiredo; José de la Fuente; Luisa De Martino; Antonella De Matteis; Guido Ry De Meyer; Angelo De Milito; Mauro De Santi; Wanderley de Souza; Vincenzo De Tata; Daniela De Zio; Jayanta Debnath; Reinhard Dechant; Jean-Paul Decuypere; Shane Deegan; Benjamin Dehay; Barbara Del Bello; Dominic P Del Re; Régis Delage-Mourroux; Lea Md Delbridge; Louise Deldicque; Elizabeth Delorme-Axford; Yizhen Deng; Joern Dengjel; Melanie Denizot; Paul Dent; Channing J Der; Vojo Deretic; Benoît Derrien; Eric Deutsch; Timothy P Devarenne; Rodney J Devenish; Sabrina Di Bartolomeo; Nicola Di Daniele; Fabio Di Domenico; Alessia Di Nardo; Simone Di Paola; Antonio Di Pietro; Livia Di Renzo; Aaron DiAntonio; Guillermo Díaz-Araya; Ines Díaz-Laviada; Maria T Diaz-Meco; Javier Diaz-Nido; Chad A Dickey; Robert C Dickson; Marc Diederich; Paul Digard; Ivan Dikic; Savithrama P Dinesh-Kumar; Chan Ding; Wen-Xing Ding; Zufeng Ding; Luciana Dini; Jörg Hw Distler; Abhinav Diwan; Mojgan Djavaheri-Mergny; Kostyantyn Dmytruk; Renwick Cj Dobson; Volker Doetsch; Karol Dokladny; Svetlana Dokudovskaya; Massimo Donadelli; X Charlie Dong; Xiaonan Dong; Zheng Dong; Terrence M Donohue; Kelly S Doran; Gabriella D'Orazi; Gerald W Dorn; Victor Dosenko; Sami Dridi; Liat Drucker; Jie Du; Li-Lin Du; Lihuan Du; André du Toit; Priyamvada Dua; Lei Duan; Pu Duann; Vikash Kumar Dubey; Michael R Duchen; Michel A Duchosal; Helene Duez; Isabelle Dugail; Verónica I Dumit; Mara C Duncan; Elaine A Dunlop; William A Dunn; Nicolas Dupont; Luc Dupuis; Raúl V Durán; Thomas M Durcan; Stéphane Duvezin-Caubet; Umamaheswar Duvvuri; Vinay Eapen; Darius Ebrahimi-Fakhari; Arnaud Echard; Leopold Eckhart; Charles L Edelstein; Aimee L Edinger; Ludwig Eichinger; Tobias Eisenberg; Avital Eisenberg-Lerner; N Tony Eissa; Wafik S El-Deiry; Victoria El-Khoury; Zvulun Elazar; Hagit Eldar-Finkelman; Chris Jh Elliott; Enzo Emanuele; Urban Emmenegger; Nikolai Engedal; Anna-Mart Engelbrecht; Simone Engelender; Jorrit M Enserink; Ralf Erdmann; Jekaterina Erenpreisa; Rajaraman Eri; Jason L Eriksen; Andreja Erman; Ricardo Escalante; Eeva-Liisa Eskelinen; Lucile Espert; Lorena Esteban-Martínez; Thomas J Evans; Mario Fabri; Gemma Fabrias; Cinzia Fabrizi; Antonio Facchiano; Nils J Færgeman; Alberto Faggioni; W Douglas Fairlie; Chunhai Fan; Daping Fan; Jie Fan; Shengyun Fang; Manolis Fanto; Alessandro Fanzani; Thomas Farkas; Mathias Faure; Francois B Favier; Howard Fearnhead; Massimo Federici; Erkang Fei; Tania C Felizardo; Hua Feng; Yibin Feng; Yuchen Feng; Thomas A Ferguson; Álvaro F Fernández; Maite G Fernandez-Barrena; Jose C Fernandez-Checa; Arsenio Fernández-López; Martin E Fernandez-Zapico; Olivier Feron; Elisabetta Ferraro; Carmen Veríssima Ferreira-Halder; Laszlo Fesus; Ralph Feuer; Fabienne C Fiesel; Eduardo C Filippi-Chiela; Giuseppe Filomeni; Gian Maria Fimia; John H Fingert; Steven Finkbeiner; Toren Finkel; Filomena Fiorito; Paul B Fisher; Marc Flajolet; Flavio Flamigni; Oliver Florey; Salvatore Florio; R Andres Floto; Marco Folini; Carlo Follo; Edward A Fon; Francesco Fornai; Franco Fortunato; Alessandro Fraldi; Rodrigo Franco; Arnaud Francois; Aurélie François; Lisa B Frankel; Iain Dc Fraser; Norbert Frey; Damien G Freyssenet; Christian Frezza; Scott L Friedman; Daniel E Frigo; Dongxu Fu; José M Fuentes; Juan Fueyo; Yoshio Fujitani; Yuuki Fujiwara; Mikihiro Fujiya; Mitsunori Fukuda; Simone Fulda; Carmela Fusco; Bozena Gabryel; Matthias Gaestel; Philippe Gailly; Malgorzata Gajewska; Sehamuddin Galadari; Gad Galili; Inmaculada Galindo; Maria F Galindo; Giovanna Galliciotti; Lorenzo Galluzzi; Luca Galluzzi; Vincent Galy; Noor Gammoh; Sam Gandy; Anand K Ganesan; Swamynathan Ganesan; Ian G Ganley; Monique Gannagé; Fen-Biao Gao; Feng Gao; Jian-Xin Gao; Lorena García Nannig; Eleonora García Véscovi; Marina Garcia-Macía; Carmen Garcia-Ruiz; Abhishek D Garg; Pramod Kumar Garg; Ricardo Gargini; Nils Christian Gassen; Damián Gatica; Evelina Gatti; Julie Gavard; Evripidis Gavathiotis; Liang Ge; Pengfei Ge; Shengfang Ge; Po-Wu Gean; Vania Gelmetti; Armando A Genazzani; Jiefei Geng; Pascal Genschik; Lisa Gerner; Jason E Gestwicki; David A Gewirtz; Saeid Ghavami; Eric Ghigo; Debabrata Ghosh; Anna Maria Giammarioli; Francesca Giampieri; Claudia Giampietri; Alexandra Giatromanolaki; Derrick J Gibbings; Lara Gibellini; Spencer B Gibson; Vanessa Ginet; Antonio Giordano; Flaviano Giorgini; Elisa Giovannetti; Stephen E Girardin; Suzana Gispert; Sandy Giuliano; Candece L Gladson; Alvaro Glavic; Martin Gleave; Nelly Godefroy; Robert M Gogal; Kuppan Gokulan; Gustavo H Goldman; Delia Goletti; Michael S Goligorsky; Aldrin V Gomes; Ligia C Gomes; Hernando Gomez; Candelaria Gomez-Manzano; Rubén Gómez-Sánchez; Dawit Ap Gonçalves; Ebru Goncu; Qingqiu Gong; Céline Gongora; Carlos B Gonzalez; Pedro Gonzalez-Alegre; Pilar Gonzalez-Cabo; Rosa Ana González-Polo; Ing Swie Goping; Carlos Gorbea; Nikolai V Gorbunov; Daphne R Goring; Adrienne M Gorman; Sharon M Gorski; Sandro Goruppi; Shino Goto-Yamada; Cecilia Gotor; Roberta A Gottlieb; Illana Gozes; Devrim Gozuacik; Yacine Graba; Martin Graef; Giovanna E Granato; Gary Dean Grant; Steven Grant; Giovanni Luca Gravina; Douglas R Green; Alexander Greenhough; Michael T Greenwood; Benedetto Grimaldi; Frédéric Gros; Charles Grose; Jean-Francois Groulx; Florian Gruber; Paolo Grumati; Tilman Grune; Jun-Lin Guan; Kun-Liang Guan; Barbara Guerra; Carlos Guillen; Kailash Gulshan; Jan Gunst; Chuanyong Guo; Lei Guo; Ming Guo; Wenjie Guo; Xu-Guang Guo; Andrea A Gust; Åsa B Gustafsson; Elaine Gutierrez; Maximiliano G Gutierrez; Ho-Shin Gwak; Albert Haas; James E Haber; Shinji Hadano; Monica Hagedorn; David R Hahn; Andrew J Halayko; Anne Hamacher-Brady; Kozo Hamada; Ahmed Hamai; Andrea Hamann; Maho Hamasaki; Isabelle Hamer; Qutayba Hamid; Ester M Hammond; Feng Han; Weidong Han; James T Handa; John A Hanover; Malene Hansen; Masaru Harada; Ljubica Harhaji-Trajkovic; J Wade Harper; Abdel Halim Harrath; Adrian L Harris; James Harris; Udo Hasler; Peter Hasselblatt; Kazuhisa Hasui; Robert G Hawley; Teresa S Hawley; Congcong He; Cynthia Y He; Fengtian He; Gu He; Rong-Rong He; Xian-Hui He; You-Wen He; Yu-Ying He; Joan K Heath; Marie-Josée Hébert; Robert A Heinzen; Gudmundur Vignir Helgason; Michael Hensel; Elizabeth P Henske; Chengtao Her; Paul K Herman; Agustín Hernández; Carlos Hernandez; Sonia Hernández-Tiedra; Claudio Hetz; P Robin Hiesinger; Katsumi Higaki; Sabine Hilfiker; Bradford G Hill; Joseph A Hill; William D Hill; Keisuke Hino; Daniel Hofius; Paul Hofman; Günter U Höglinger; Jörg Höhfeld; Marina K Holz; Yonggeun Hong; David A Hood; Jeroen Jm Hoozemans; Thorsten Hoppe; Chin Hsu; Chin-Yuan Hsu; Li-Chung Hsu; Dong Hu; Guochang Hu; Hong-Ming Hu; Hongbo Hu; Ming Chang Hu; Yu-Chen Hu; Zhuo-Wei Hu; Fang Hua; Ya Hua; Canhua Huang; Huey-Lan Huang; Kuo-How Huang; Kuo-Yang Huang; Shile Huang; Shiqian Huang; Wei-Pang Huang; Yi-Ran Huang; Yong Huang; Yunfei Huang; Tobias B Huber; Patricia Huebbe; Won-Ki Huh; Juha J Hulmi; Gang Min Hur; James H Hurley; Zvenyslava Husak; Sabah Na Hussain; Salik Hussain; Jung Jin Hwang; Seungmin Hwang; Thomas Is Hwang; Atsuhiro Ichihara; Yuzuru Imai; Carol Imbriano; Megumi Inomata; Takeshi Into; Valentina Iovane; Juan L Iovanna; Renato V Iozzo; Nancy Y Ip; Javier E Irazoqui; Pablo Iribarren; Yoshitaka Isaka; Aleksandra J Isakovic; Harry Ischiropoulos; Jeffrey S Isenberg; Mohammad Ishaq; Hiroyuki Ishida; Isao Ishii; Jane E Ishmael; Ciro Isidoro; Ken-Ichi Isobe; Erika Isono; Shohreh Issazadeh-Navikas; Koji Itahana; Eisuke Itakura; Andrei I Ivanov; Anand Krishnan V Iyer; José M Izquierdo; Yotaro Izumi; Valentina Izzo; Marja Jäättelä; Nadia Jaber; Daniel John Jackson; William T Jackson; Tony George Jacob; Thomas S Jacques; Chinnaswamy Jagannath; Ashish Jain; Nihar Ranjan Jana; Byoung Kuk Jang; Alkesh Jani; Bassam Janji; Paulo Roberto Jannig; Patric J Jansson; Steve Jean; Marina Jendrach; Ju-Hong Jeon; Niels Jessen; Eui-Bae Jeung; Kailiang Jia; Lijun Jia; Hong Jiang; Hongchi Jiang; Liwen Jiang; Teng Jiang; Xiaoyan Jiang; Xuejun Jiang; Xuejun Jiang; Ying Jiang; Yongjun Jiang; Alberto Jiménez; Cheng Jin; Hongchuan Jin; Lei Jin; Meiyan Jin; Shengkan Jin; Umesh Kumar Jinwal; Eun-Kyeong Jo; Terje Johansen; Daniel E Johnson; Gail Vw Johnson; James D Johnson; Eric Jonasch; Chris Jones; Leo Ab Joosten; Joaquin Jordan; Anna-Maria Joseph; Bertrand Joseph; Annie M Joubert; Dianwen Ju; Jingfang Ju; Hsueh-Fen Juan; Katrin Juenemann; Gábor Juhász; Hye Seung Jung; Jae U Jung; Yong-Keun Jung; Heinz Jungbluth; Matthew J Justice; Barry Jutten; Nadeem O Kaakoush; Kai Kaarniranta; Allen Kaasik; Tomohiro Kabuta; Bertrand Kaeffer; Katarina Kågedal; Alon Kahana; Shingo Kajimura; Or Kakhlon; Manjula Kalia; Dhan V Kalvakolanu; Yoshiaki Kamada; Konstantinos Kambas; Vitaliy O Kaminskyy; Harm H Kampinga; Mustapha Kandouz; Chanhee Kang; Rui Kang; Tae-Cheon Kang; Tomotake Kanki; Thirumala-Devi Kanneganti; Haruo Kanno; Anumantha G Kanthasamy; Marc Kantorow; Maria Kaparakis-Liaskos; Orsolya Kapuy; Vassiliki Karantza; Md Razaul Karim; Parimal Karmakar; Arthur Kaser; Susmita Kaushik; Thomas Kawula; A Murat Kaynar; Po-Yuan Ke; Zun-Ji Ke; John H Kehrl; Kate E Keller; Jongsook Kim Kemper; Anne K Kenworthy; Oliver Kepp; Andreas Kern; Santosh Kesari; David Kessel; Robin Ketteler; Isis do Carmo Kettelhut; Bilon Khambu; Muzamil Majid Khan; Vinoth Km Khandelwal; Sangeeta Khare; Juliann G Kiang; Amy A Kiger; Akio Kihara; Arianna L Kim; Cheol Hyeon Kim; Deok Ryong Kim; Do-Hyung Kim; Eung Kweon Kim; Hye Young Kim; Hyung-Ryong Kim; Jae-Sung Kim; Jeong Hun Kim; Jin Cheon Kim; Jin Hyoung Kim; Kwang Woon Kim; Michael D Kim; Moon-Moo Kim; Peter K Kim; Seong Who Kim; Soo-Youl Kim; Yong-Sun Kim; Yonghyun Kim; Adi Kimchi; Alec C Kimmelman; Tomonori Kimura; Jason S King; Karla Kirkegaard; Vladimir Kirkin; Lorrie A Kirshenbaum; Shuji Kishi; Yasuo Kitajima; Katsuhiko Kitamoto; Yasushi Kitaoka; Kaio Kitazato; Rudolf A Kley; Walter T Klimecki; Michael Klinkenberg; Jochen Klucken; Helene Knævelsrud; Erwin Knecht; Laura Knuppertz; Jiunn-Liang Ko; Satoru Kobayashi; Jan C Koch; Christelle Koechlin-Ramonatxo; Ulrich Koenig; Young Ho Koh; Katja Köhler; Sepp D Kohlwein; Masato Koike; Masaaki Komatsu; Eiki Kominami; Dexin Kong; Hee Jeong Kong; Eumorphia G Konstantakou; Benjamin T Kopp; Tamas Korcsmaros; Laura Korhonen; Viktor I Korolchuk; Nadya V Koshkina; Yanjun Kou; Michael I Koukourakis; Constantinos Koumenis; Attila L Kovács; Tibor Kovács; Werner J Kovacs; Daisuke Koya; Claudine Kraft; Dimitri Krainc; Helmut Kramer; Tamara Kravic-Stevovic; Wilhelm Krek; Carole Kretz-Remy; Roswitha Krick; Malathi Krishnamurthy; Janos Kriston-Vizi; Guido Kroemer; Michael C Kruer; Rejko Kruger; Nicholas T Ktistakis; Kazuyuki Kuchitsu; Christian Kuhn; Addanki Pratap Kumar; Anuj Kumar; Ashok Kumar; Deepak Kumar; Dhiraj Kumar; Rakesh Kumar; Sharad Kumar; Mondira Kundu; Hsing-Jien Kung; Atsushi Kuno; Sheng-Han Kuo; Jeff Kuret; Tino Kurz; Terry Kwok; Taeg Kyu Kwon; Yong Tae Kwon; Irene Kyrmizi; Albert R La Spada; Frank Lafont; Tim Lahm; Aparna Lakkaraju; Truong Lam; Trond Lamark; Steve Lancel; Terry H Landowski; Darius J R Lane; Jon D Lane; Cinzia Lanzi; Pierre Lapaquette; Louis R Lapierre; Jocelyn Laporte; Johanna Laukkarinen; Gordon W Laurie; Sergio Lavandero; Lena Lavie; Matthew J LaVoie; Betty Yuen Kwan Law; Helen Ka-Wai Law; Kelsey B Law; Robert Layfield; Pedro A Lazo; Laurent Le Cam; Karine G Le Roch; Hervé Le Stunff; Vijittra Leardkamolkarn; Marc Lecuit; Byung-Hoon Lee; Che-Hsin Lee; Erinna F Lee; Gyun Min Lee; He-Jin Lee; Hsinyu Lee; Jae Keun Lee; Jongdae Lee; Ju-Hyun Lee; Jun Hee Lee; Michael Lee; Myung-Shik Lee; Patty J Lee; Sam W Lee; Seung-Jae Lee; Shiow-Ju Lee; Stella Y Lee; Sug Hyung Lee; Sung Sik Lee; Sung-Joon Lee; Sunhee Lee; Ying-Ray Lee; Yong J Lee; Young H Lee; Christiaan Leeuwenburgh; Sylvain Lefort; Renaud Legouis; Jinzhi Lei; Qun-Ying Lei; David A Leib; Gil Leibowitz; Istvan Lekli; Stéphane D Lemaire; John J Lemasters; Marius K Lemberg; Antoinette Lemoine; Shuilong Leng; Guido Lenz; Paola Lenzi; Lilach O Lerman; Daniele Lettieri Barbato; Julia I-Ju Leu; Hing Y Leung; Beth Levine; Patrick A Lewis; Frank Lezoualc'h; Chi Li; Faqiang Li; Feng-Jun Li; Jun Li; Ke Li; Lian Li; Min Li; Min Li; Qiang Li; Rui Li; Sheng Li; Wei Li; Wei Li; Xiaotao Li; Yumin Li; Jiqin Lian; Chengyu Liang; Qiangrong Liang; Yulin Liao; Joana Liberal; Pawel P Liberski; Pearl Lie; Andrew P Lieberman; Hyunjung Jade Lim; Kah-Leong Lim; Kyu Lim; Raquel T Lima; Chang-Shen Lin; Chiou-Feng Lin; Fang Lin; Fangming Lin; Fu-Cheng Lin; Kui Lin; Kwang-Huei Lin; Pei-Hui Lin; Tianwei Lin; Wan-Wan Lin; Yee-Shin Lin; Yong Lin; Rafael Linden; Dan Lindholm; Lisa M Lindqvist; Paul Lingor; Andreas Linkermann; Lance A Liotta; Marta M Lipinski; Vitor A Lira; Michael P Lisanti; Paloma B Liton; Bo Liu; Chong Liu; Chun-Feng Liu; Fei Liu; Hung-Jen Liu; Jianxun Liu; Jing-Jing Liu; Jing-Lan Liu; Ke Liu; Leyuan Liu; Liang Liu; Quentin Liu; Rong-Yu Liu; Shiming Liu; Shuwen Liu; Wei Liu; Xian-De Liu; Xiangguo Liu; Xiao-Hong Liu; Xinfeng Liu; Xu Liu; Xueqin Liu; Yang Liu; Yule Liu; Zexian Liu; Zhe Liu; Juan P Liuzzi; Gérard Lizard; Mila Ljujic; Irfan J Lodhi; Susan E Logue; Bal L Lokeshwar; Yun Chau Long; Sagar Lonial; Benjamin Loos; Carlos López-Otín; Cristina López-Vicario; Mar Lorente; Philip L Lorenzi; Péter Lõrincz; Marek Los; Michael T Lotze; Penny E Lovat; Binfeng Lu; Bo Lu; Jiahong Lu; Qing Lu; She-Min Lu; Shuyan Lu; Yingying Lu; Frédéric Luciano; Shirley Luckhart; John Milton Lucocq; Paula Ludovico; Aurelia Lugea; Nicholas W Lukacs; Julian J Lum; Anders H Lund; Honglin Luo; Jia Luo; Shouqing Luo; Claudio Luparello; Timothy Lyons; Jianjie Ma; Yi Ma; Yong Ma; Zhenyi Ma; Juliano Machado; Glaucia M Machado-Santelli; Fernando Macian; Gustavo C MacIntosh; Jeffrey P MacKeigan; Kay F Macleod; John D MacMicking; Lee Ann MacMillan-Crow; Frank Madeo; Muniswamy Madesh; Julio Madrigal-Matute; Akiko Maeda; Tatsuya Maeda; Gustavo Maegawa; Emilia Maellaro; Hannelore Maes; Marta Magariños; Kenneth Maiese; Tapas K Maiti; Luigi Maiuri; Maria Chiara Maiuri; Carl G Maki; Roland Malli; Walter Malorni; Alina Maloyan; Fathia Mami-Chouaib; Na Man; Joseph D Mancias; Eva-Maria Mandelkow; Michael A Mandell; Angelo A Manfredi; Serge N Manié; Claudia Manzoni; Kai Mao; Zixu Mao; Zong-Wan Mao; Philippe Marambaud; Anna Maria Marconi; Zvonimir Marelja; Gabriella Marfe; Marta Margeta; Eva Margittai; Muriel Mari; Francesca V Mariani; Concepcio Marin; Sara Marinelli; Guillermo Mariño; Ivanka Markovic; Rebecca Marquez; Alberto M Martelli; Sascha Martens; Katie R Martin; Seamus J Martin; Shaun Martin; Miguel A Martin-Acebes; Paloma Martín-Sanz; Camille Martinand-Mari; Wim Martinet; Jennifer Martinez; Nuria Martinez-Lopez; Ubaldo Martinez-Outschoorn; Moisés Martínez-Velázquez; Marta Martinez-Vicente; Waleska Kerllen Martins; Hirosato Mashima; James A Mastrianni; Giuseppe Matarese; Paola Matarrese; Roberto Mateo; Satoaki Matoba; Naomichi Matsumoto; Takehiko Matsushita; Akira Matsuura; Takeshi Matsuzawa; Mark P Mattson; Soledad Matus; Norma Maugeri; Caroline Mauvezin; Andreas Mayer; Dusica Maysinger; Guillermo D Mazzolini; Mary Kate McBrayer; Kimberly McCall; Craig McCormick; Gerald M McInerney; Skye C McIver; Sharon McKenna; John J McMahon; Iain A McNeish; Fatima Mechta-Grigoriou; Jan Paul Medema; Diego L Medina; Klara Megyeri; Maryam Mehrpour; Jawahar L Mehta; Yide Mei; Ute-Christiane Meier; Alfred J Meijer; Alicia Meléndez; Gerry Melino; Sonia Melino; Edesio Jose Tenorio de Melo; Maria A Mena; Marc D Meneghini; Javier A Menendez; Regina Menezes; Liesu Meng; Ling-Hua Meng; Songshu Meng; Rossella Menghini; A Sue Menko; Rubem Fs Menna-Barreto; Manoj B Menon; Marco A Meraz-Ríos; Giuseppe Merla; Luciano Merlini; Angelica M Merlot; Andreas Meryk; Stefania Meschini; Joel N Meyer; Man-Tian Mi; Chao-Yu Miao; Lucia Micale; Simon Michaeli; Carine Michiels; Anna Rita Migliaccio; Anastasia Susie Mihailidou; Dalibor Mijaljica; Katsuhiko Mikoshiba; Enrico Milan; Leonor Miller-Fleming; Gordon B Mills; Ian G Mills; Georgia Minakaki; Berge A Minassian; Xiu-Fen Ming; Farida Minibayeva; Elena A Minina; Justine D Mintern; Saverio Minucci; Antonio Miranda-Vizuete; Claire H Mitchell; Shigeki Miyamoto; Keisuke Miyazawa; Noboru Mizushima; Katarzyna Mnich; Baharia Mograbi; Simin Mohseni; Luis Ferreira Moita; Marco Molinari; Maurizio Molinari; Andreas Buch Møller; Bertrand Mollereau; Faustino Mollinedo; Marco Mongillo; Martha M Monick; Serena Montagnaro; Craig Montell; Darren J Moore; Michael N Moore; Rodrigo Mora-Rodriguez; Paula I Moreira; Etienne Morel; Maria Beatrice Morelli; Sandra Moreno; Michael J Morgan; Arnaud Moris; Yuji Moriyasu; Janna L Morrison; Lynda A Morrison; Eugenia Morselli; Jorge Moscat; Pope L Moseley; Serge Mostowy; Elisa Motori; Denis Mottet; Jeremy C Mottram; Charbel E-H Moussa; Vassiliki E Mpakou; Hasan Mukhtar; Jean M Mulcahy Levy; Sylviane Muller; Raquel Muñoz-Moreno; Cristina Muñoz-Pinedo; Christian Münz; Maureen E Murphy; James T Murray; Aditya Murthy; Indira U Mysorekar; Ivan R Nabi; Massimo Nabissi; Gustavo A Nader; Yukitoshi Nagahara; Yoshitaka Nagai; Kazuhiro Nagata; Anika Nagelkerke; Péter Nagy; Samisubbu R Naidu; Sreejayan Nair; Hiroyasu Nakano; Hitoshi Nakatogawa; Meera Nanjundan; Gennaro Napolitano; Naweed I Naqvi; Roberta Nardacci; Derek P Narendra; Masashi Narita; Anna Chiara Nascimbeni; Ramesh Natarajan; Luiz C Navegantes; Steffan T Nawrocki; Taras Y Nazarko; Volodymyr Y Nazarko; Thomas Neill; Luca M Neri; Mihai G Netea; Romana T Netea-Maier; Bruno M Neves; Paul A Ney; Ioannis P Nezis; Hang Tt Nguyen; Huu Phuc Nguyen; Anne-Sophie Nicot; Hilde Nilsen; Per Nilsson; Mikio Nishimura; Ichizo Nishino; Mireia Niso-Santano; Hua Niu; Ralph A Nixon; Vincent Co Njar; Takeshi Noda; Angelika A Noegel; Elsie Magdalena Nolte; Erik Norberg; Koenraad K Norga; Sakineh Kazemi Noureini; Shoji Notomi; Lucia Notterpek; Karin Nowikovsky; Nobuyuki Nukina; Thorsten Nürnberger; Valerie B O'Donnell; Tracey O'Donovan; Peter J O'Dwyer; Ina Oehme; Clara L Oeste; Michinaga Ogawa; Besim Ogretmen; Yuji Ogura; Young J Oh; Masaki Ohmuraya; Takayuki Ohshima; Rani Ojha; Koji Okamoto; Toshiro Okazaki; F Javier Oliver; Karin Ollinger; Stefan Olsson; Daniel P Orban; Paulina Ordonez; Idil Orhon; Laszlo Orosz; Eyleen J O'Rourke; Helena Orozco; Angel L Ortega; Elena Ortona; Laura D Osellame; Junko Oshima; Shigeru Oshima; Heinz D Osiewacz; Takanobu Otomo; Kinya Otsu; Jing-Hsiung James Ou; Tiago F Outeiro; Dong-Yun Ouyang; Hongjiao Ouyang; Michael Overholtzer; Michelle A Ozbun; P Hande Ozdinler; Bulent Ozpolat; Consiglia Pacelli; Paolo Paganetti; Guylène Page; Gilles Pages; Ugo Pagnini; Beata Pajak; Stephen C Pak; Karolina Pakos-Zebrucka; Nazzy Pakpour; Zdena Palková; Francesca Palladino; Kathrin Pallauf; Nicolas Pallet; Marta Palmieri; Søren R Paludan; Camilla Palumbo; Silvia Palumbo; Olatz Pampliega; Hongming Pan; Wei Pan; Theocharis Panaretakis; Aseem Pandey; Areti Pantazopoulou; Zuzana Papackova; Daniela L Papademetrio; Issidora Papassideri; Alessio Papini; Nirmala Parajuli; Julian Pardo; Vrajesh V Parekh; Giancarlo Parenti; Jong-In Park; Junsoo Park; Ohkmae K Park; Roy Parker; Rosanna Parlato; Jan B Parys; Katherine R Parzych; Jean-Max Pasquet; Benoit Pasquier; Kishore Bs Pasumarthi; Daniel Patschan; Cam Patterson; Sophie Pattingre; Scott Pattison; Arnim Pause; Hermann Pavenstädt; Flaminia Pavone; Zully Pedrozo; Fernando J Peña; Miguel A Peñalva; Mario Pende; Jianxin Peng; Fabio Penna; Josef M Penninger; Anna Pensalfini; Salvatore Pepe; Gustavo Js Pereira; Paulo C Pereira; Verónica Pérez-de la Cruz; María Esther Pérez-Pérez; Diego Pérez-Rodríguez; Dolores Pérez-Sala; Celine Perier; Andras Perl; David H Perlmutter; Ida Perrotta; Shazib Pervaiz; Maija Pesonen; Jeffrey E Pessin; Godefridus J Peters; Morten Petersen; Irina Petrache; Basil J Petrof; Goran Petrovski; James M Phang; Mauro Piacentini; Marina Pierdominici; Philippe Pierre; Valérie Pierrefite-Carle; Federico Pietrocola; Felipe X Pimentel-Muiños; Mario Pinar; Benjamin Pineda; Ronit Pinkas-Kramarski; Marcello Pinti; Paolo Pinton; Bilal Piperdi; James M Piret; Leonidas C Platanias; Harald W Platta; Edward D Plowey; Stefanie Pöggeler; Marc Poirot; Peter Polčic; Angelo Poletti; Audrey H Poon; Hana Popelka; Blagovesta Popova; Izabela Poprawa; Shibu M Poulose; Joanna Poulton; Scott K Powers; Ted Powers; Mercedes Pozuelo-Rubio; Krisna Prak; Reinhild Prange; Mark Prescott; Muriel Priault; Sharon Prince; Richard L Proia; Tassula Proikas-Cezanne; Holger Prokisch; Vasilis J Promponas; Karin Przyklenk; Rosa Puertollano; Subbiah Pugazhenthi; Luigi Puglielli; Aurora Pujol; Julien Puyal; Dohun Pyeon; Xin Qi; Wen-Bin Qian; Zheng-Hong Qin; Yu Qiu; Ziwei Qu; Joe Quadrilatero; Frederick Quinn; Nina Raben; Hannah Rabinowich; Flavia Radogna; Michael J Ragusa; Mohamed Rahmani; Komal Raina; Sasanka Ramanadham; Rajagopal Ramesh; Abdelhaq Rami; Sarron Randall-Demllo; Felix Randow; Hai Rao; V Ashutosh Rao; Blake B Rasmussen; Tobias M Rasse; Edward A Ratovitski; Pierre-Emmanuel Rautou; Swapan K Ray; Babak Razani; Bruce H Reed; Fulvio Reggiori; Markus Rehm; Andreas S Reichert; Theo Rein; David J Reiner; Eric Reits; Jun Ren; Xingcong Ren; Maurizio Renna; Jane Eb Reusch; Jose L Revuelta; Leticia Reyes; Alireza R Rezaie; Robert I Richards; Des R Richardson; Clémence Richetta; Michael A Riehle; Bertrand H Rihn; Yasuko Rikihisa; Brigit E Riley; Gerald Rimbach; Maria Rita Rippo; Konstantinos Ritis; Federica Rizzi; Elizete Rizzo; Peter J Roach; Jeffrey Robbins; Michel Roberge; Gabriela Roca; Maria Carmela Roccheri; Sonia Rocha; Cecilia Mp Rodrigues; Clara I Rodríguez; Santiago Rodriguez de Cordoba; Natalia Rodriguez-Muela; Jeroen Roelofs; Vladimir V Rogov; Troy T Rohn; Bärbel Rohrer; Davide Romanelli; Luigina Romani; Patricia Silvia Romano; M Isabel G Roncero; Jose Luis Rosa; Alicia Rosello; Kirill V Rosen; Philip Rosenstiel; Magdalena Rost-Roszkowska; Kevin A Roth; Gael Roué; Mustapha Rouis; Kasper M Rouschop; Daniel T Ruan; Diego Ruano; David C Rubinsztein; Edmund B Rucker; Assaf Rudich; Emil Rudolf; Ruediger Rudolf; Markus A Ruegg; Carmen Ruiz-Roldan; Avnika Ashok Ruparelia; Paola Rusmini; David W Russ; Gian Luigi Russo; Giuseppe Russo; Rossella Russo; Tor Erik Rusten; Victoria Ryabovol; Kevin M Ryan; Stefan W Ryter; David M Sabatini; Michael Sacher; Carsten Sachse; Michael N Sack; Junichi Sadoshima; Paul Saftig; Ronit Sagi-Eisenberg; Sumit Sahni; Pothana Saikumar; Tsunenori Saito; Tatsuya Saitoh; Koichi Sakakura; Machiko Sakoh-Nakatogawa; Yasuhito Sakuraba; María Salazar-Roa; Paolo Salomoni; Ashok K Saluja; Paul M Salvaterra; Rosa Salvioli; Afshin Samali; Anthony Mj Sanchez; José A Sánchez-Alcázar; Ricardo Sanchez-Prieto; Marco Sandri; Miguel A Sanjuan; Stefano Santaguida; Laura Santambrogio; Giorgio Santoni; Claudia Nunes Dos Santos; Shweta Saran; Marco Sardiello; Graeme Sargent; Pallabi Sarkar; Sovan Sarkar; Maria Rosa Sarrias; Minnie M Sarwal; Chihiro Sasakawa; Motoko Sasaki; Miklos Sass; Ken Sato; Miyuki Sato; Joseph Satriano; Niramol Savaraj; Svetlana Saveljeva; Liliana Schaefer; Ulrich E Schaible; Michael Scharl; Hermann M Schatzl; Randy Schekman; Wiep Scheper; Alfonso Schiavi; Hyman M Schipper; Hana Schmeisser; Jens Schmidt; Ingo Schmitz; Bianca E Schneider; E Marion Schneider; Jaime L Schneider; Eric A Schon; Miriam J Schönenberger; Axel H Schönthal; Daniel F Schorderet; Bernd Schröder; Sebastian Schuck; Ryan J Schulze; Melanie Schwarten; Thomas L Schwarz; Sebastiano Sciarretta; Kathleen Scotto; A Ivana Scovassi; Robert A Screaton; Mark Screen; Hugo Seca; Simon Sedej; Laura Segatori; Nava Segev; Per O Seglen; Jose M Seguí-Simarro; Juan Segura-Aguilar; Ekihiro Seki; Christian Sell; Iban Seiliez; Clay F Semenkovich; Gregg L Semenza; Utpal Sen; Andreas L Serra; Ana Serrano-Puebla; Hiromi Sesaki; Takao Setoguchi; Carmine Settembre; John J Shacka; Ayesha N Shajahan-Haq; Irving M Shapiro; Shweta Sharma; Hua She; C-K James Shen; Chiung-Chyi Shen; Han-Ming Shen; Sanbing Shen; Weili Shen; Rui Sheng; Xianyong Sheng; Zu-Hang Sheng; Trevor G Shepherd; Junyan Shi; Qiang Shi; Qinghua Shi; Yuguang Shi; Shusaku Shibutani; Kenichi Shibuya; Yoshihiro Shidoji; Jeng-Jer Shieh; Chwen-Ming Shih; Yohta Shimada; Shigeomi Shimizu; Dong Wook Shin; Mari L Shinohara; Michiko Shintani; Takahiro Shintani; Tetsuo Shioi; Ken Shirabe; Ronit Shiri-Sverdlov; Orian Shirihai; Gordon C Shore; Chih-Wen Shu; Deepak Shukla; Andriy A Sibirny; Valentina Sica; Christina J Sigurdson; Einar M Sigurdsson; Puran Singh Sijwali; Beata Sikorska; Wilian A Silveira; Sandrine Silvente-Poirot; Gary A Silverman; Jan Simak; Thomas Simmet; Anna Katharina Simon; Hans-Uwe Simon; Cristiano Simone; Matias Simons; Anne Simonsen; Rajat Singh; Shivendra V Singh; Shrawan K Singh; Debasish Sinha; Sangita Sinha; Frank A Sinicrope; Agnieszka Sirko; Kapil Sirohi; Balindiwe Jn Sishi; Annie Sittler; Parco M Siu; Efthimios Sivridis; Anna Skwarska; Ruth Slack; Iva Slaninová; Nikolai Slavov; Soraya S Smaili; Keiran Sm Smalley; Duncan R Smith; Stefaan J Soenen; Scott A Soleimanpour; Anita Solhaug; Kumaravel Somasundaram; Jin H Son; Avinash Sonawane; Chunjuan Song; Fuyong Song; Hyun Kyu Song; Ju-Xian Song; Wei Song; Kai Y Soo; Anil K Sood; Tuck Wah Soong; Virawudh Soontornniyomkij; Maurizio Sorice; Federica Sotgia; David R Soto-Pantoja; Areechun Sotthibundhu; Maria João Sousa; Herman P Spaink; Paul N Span; Anne Spang; Janet D Sparks; Peter G Speck; Stephen A Spector; Claudia D Spies; Wolfdieter Springer; Daret St Clair; Alessandra Stacchiotti; Bart Staels; Michael T Stang; Daniel T Starczynowski; Petro Starokadomskyy; Clemens Steegborn; John W Steele; Leonidas Stefanis; Joan Steffan; Christine M Stellrecht; Harald Stenmark; Tomasz M Stepkowski; Stęphan T Stern; Craig Stevens; Brent R Stockwell; Veronika Stoka; Zuzana Storchova; Björn Stork; Vassilis Stratoulias; Dimitrios J Stravopodis; Pavel Strnad; Anne Marie Strohecker; Anna-Lena Ström; Per Stromhaug; Jiri Stulik; Yu-Xiong Su; Zhaoliang Su; Carlos S Subauste; Srinivasa Subramaniam; Carolyn M Sue; Sang Won Suh; Xinbing Sui; Supawadee Sukseree; David Sulzer; Fang-Lin Sun; Jiaren Sun; Jun Sun; Shi-Yong Sun; Yang Sun; Yi Sun; Yingjie Sun; Vinod Sundaramoorthy; Joseph Sung; Hidekazu Suzuki; Kuninori Suzuki; Naoki Suzuki; Tadashi Suzuki; Yuichiro J Suzuki; Michele S Swanson; Charles Swanton; Karl Swärd; Ghanshyam Swarup; Sean T Sweeney; Paul W Sylvester; Zsuzsanna Szatmari; Eva Szegezdi; Peter W Szlosarek; Heinrich Taegtmeyer; Marco Tafani; Emmanuel Taillebourg; Stephen Wg Tait; Krisztina Takacs-Vellai; Yoshinori Takahashi; Szabolcs Takáts; Genzou Takemura; Nagio Takigawa; Nicholas J Talbot; Elena Tamagno; Jerome Tamburini; Cai-Ping Tan; Lan Tan; Mei Lan Tan; Ming Tan; Yee-Joo Tan; Keiji Tanaka; Masaki Tanaka; Daolin Tang; Dingzhong Tang; Guomei Tang; Isei Tanida; Kunikazu Tanji; Bakhos A Tannous; Jose A Tapia; Inmaculada Tasset-Cuevas; Marc Tatar; Iman Tavassoly; Nektarios Tavernarakis; Allen Taylor; Graham S Taylor; Gregory A Taylor; J Paul Taylor; Mark J Taylor; Elena V Tchetina; Andrew R Tee; Fatima Teixeira-Clerc; Sucheta Telang; Tewin Tencomnao; Ba-Bie Teng; Ru-Jeng Teng; Faraj Terro; Gianluca Tettamanti; Arianne L Theiss; Anne E Theron; Kelly Jean Thomas; Marcos P Thomé; Paul G Thomes; Andrew Thorburn; Jeremy Thorner; Thomas Thum; Michael Thumm; Teresa Lm Thurston; Ling Tian; Andreas Till; Jenny Pan-Yun Ting; Vladimir I Titorenko; Lilach Toker; Stefano Toldo; Sharon A Tooze; Ivan Topisirovic; Maria Lyngaas Torgersen; Liliana Torosantucci; Alicia Torriglia; Maria Rosaria Torrisi; Cathy Tournier; Roberto Towns; Vladimir Trajkovic; Leonardo H Travassos; Gemma Triola; Durga Nand Tripathi; Daniela Trisciuoglio; Rodrigo Troncoso; Ioannis P Trougakos; Anita C Truttmann; Kuen-Jer Tsai; Mario P Tschan; Yi-Hsin Tseng; Takayuki Tsukuba; Allan Tsung; Andrey S Tsvetkov; Shuiping Tu; Hsing-Yu Tuan; Marco Tucci; David A Tumbarello; Boris Turk; Vito Turk; Robin Fb Turner; Anders A Tveita; Suresh C Tyagi; Makoto Ubukata; Yasuo Uchiyama; Andrej Udelnow; Takashi Ueno; Midori Umekawa; Rika Umemiya-Shirafuji; Benjamin R Underwood; Christian Ungermann; Rodrigo P Ureshino; Ryo Ushioda; Vladimir N Uversky; Néstor L Uzcátegui; Thomas Vaccari; Maria I Vaccaro; Libuše Váchová; Helin Vakifahmetoglu-Norberg; Rut Valdor; Enza Maria Valente; Francois Vallette; Angela M Valverde; Greet Van den Berghe; Ludo Van Den Bosch; Gijs R van den Brink; F Gisou van der Goot; Ida J van der Klei; Luc Jw van der Laan; Wouter G van Doorn; Marjolein van Egmond; Kenneth L van Golen; Luc Van Kaer; Menno van Lookeren Campagne; Peter Vandenabeele; Wim Vandenberghe; Ilse Vanhorebeek; Isabel Varela-Nieto; M Helena Vasconcelos; Radovan Vasko; Demetrios G Vavvas; Ignacio Vega-Naredo; Guillermo Velasco; Athanassios D Velentzas; Panagiotis D Velentzas; Tibor Vellai; Edo Vellenga; Mikkel Holm Vendelbo; Kartik Venkatachalam; Natascia Ventura; Salvador Ventura; Patrícia St Veras; Mireille Verdier; Beata G Vertessy; Andrea Viale; Michel Vidal; Helena L A Vieira; Richard D Vierstra; Nadarajah Vigneswaran; Neeraj Vij; Miquel Vila; Margarita Villar; Victor H Villar; Joan Villarroya; Cécile Vindis; Giampietro Viola; Maria Teresa Viscomi; Giovanni Vitale; Dan T Vogl; Olga V Voitsekhovskaja; Clarissa von Haefen; Karin von Schwarzenberg; Daniel E Voth; Valérie Vouret-Craviari; Kristina Vuori; Jatin M Vyas; Christian Waeber; Cheryl Lyn Walker; Mark J Walker; Jochen Walter; Lei Wan; Xiangbo Wan; Bo Wang; Caihong Wang; Chao-Yung Wang; Chengshu Wang; Chenran Wang; Chuangui Wang; Dong Wang; Fen Wang; Fuxin Wang; Guanghui Wang; Hai-Jie Wang; Haichao Wang; Hong-Gang Wang; Hongmin Wang; Horng-Dar Wang; Jing Wang; Junjun Wang; Mei Wang; Mei-Qing Wang; Pei-Yu Wang; Peng Wang; Richard C Wang; Shuo Wang; Ting-Fang Wang; Xian Wang; Xiao-Jia Wang; Xiao-Wei Wang; Xin Wang; Xuejun Wang; Yan Wang; Yanming Wang; Ying Wang; Ying-Jan Wang; Yipeng Wang; Yu Wang; Yu Tian Wang; Yuqing Wang; Zhi-Nong Wang; Pablo Wappner; Carl Ward; Diane McVey Ward; Gary Warnes; Hirotaka Watada; Yoshihisa Watanabe; Kei Watase; Timothy E Weaver; Colin D Weekes; Jiwu Wei; Thomas Weide; Conrad C Weihl; Günther Weindl; Simone Nardin Weis; Longping Wen; Xin Wen; Yunfei Wen; Benedikt Westermann; Cornelia M Weyand; Anthony R White; Eileen White; J Lindsay Whitton; Alexander J Whitworth; Joëlle Wiels; Franziska Wild; Manon E Wildenberg; Tom Wileman; Deepti Srinivas Wilkinson; Simon Wilkinson; Dieter Willbold; Chris Williams; Katherine Williams; Peter R Williamson; Konstanze F Winklhofer; Steven S Witkin; Stephanie E Wohlgemuth; Thomas Wollert; Ernst J Wolvetang; Esther Wong; G William Wong; Richard W Wong; Vincent Kam Wai Wong; Elizabeth A Woodcock; Karen L Wright; Chunlai Wu; Defeng Wu; Gen Sheng Wu; Jian Wu; Junfang Wu; Mian Wu; Min Wu; Shengzhou Wu; William Kk Wu; Yaohua Wu; Zhenlong Wu; Cristina Pr Xavier; Ramnik J Xavier; Gui-Xian Xia; Tian Xia; Weiliang Xia; Yong Xia; Hengyi Xiao; Jian Xiao; Shi Xiao; Wuhan Xiao; Chuan-Ming Xie; Zhiping Xie; Zhonglin Xie; Maria Xilouri; Yuyan Xiong; Chuanshan Xu; Congfeng Xu; Feng Xu; Haoxing Xu; Hongwei Xu; Jian Xu; Jianzhen Xu; Jinxian Xu; Liang Xu; Xiaolei Xu; Yangqing Xu; Ye Xu; Zhi-Xiang Xu; Ziheng Xu; Yu Xue; Takahiro Yamada; Ai Yamamoto; Koji Yamanaka; Shunhei Yamashina; Shigeko Yamashiro; Bing Yan; Bo Yan; Xianghua Yan; Zhen Yan; Yasuo Yanagi; Dun-Sheng Yang; Jin-Ming Yang; Liu Yang; Minghua Yang; Pei-Ming Yang; Peixin Yang; Qian Yang; Wannian Yang; Wei Yuan Yang; Xuesong Yang; Yi Yang; Ying Yang; Zhifen Yang; Zhihong Yang; Meng-Chao Yao; Pamela J Yao; Xiaofeng Yao; Zhenyu Yao; Zhiyuan Yao; Linda S Yasui; Mingxiang Ye; Barry Yedvobnick; Behzad Yeganeh; Elizabeth S Yeh; Patricia L Yeyati; Fan Yi; Long Yi; Xiao-Ming Yin; Calvin K Yip; Yeong-Min Yoo; Young Hyun Yoo; Seung-Yong Yoon; Ken-Ichi Yoshida; Tamotsu Yoshimori; Ken H Young; Huixin Yu; Jane J Yu; Jin-Tai Yu; Jun Yu; Li Yu; W Haung Yu; Xiao-Fang Yu; Zhengping Yu; Junying Yuan; Zhi-Min Yuan; Beatrice Yjt Yue; Jianbo Yue; Zhenyu Yue; David N Zacks; Eldad Zacksenhaus; Nadia Zaffaroni; Tania Zaglia; Zahra Zakeri; Vincent Zecchini; Jinsheng Zeng; Min Zeng; Qi Zeng; Antonis S Zervos; Donna D Zhang; Fan Zhang; Guo Zhang; Guo-Chang Zhang; Hao Zhang; Hong Zhang; Hong Zhang; Hongbing Zhang; Jian Zhang; Jian Zhang; Jiangwei Zhang; Jianhua Zhang; Jing-Pu Zhang; Li Zhang; Lin Zhang; Lin Zhang; Long Zhang; Ming-Yong Zhang; Xiangnan Zhang; Xu Dong Zhang; Yan Zhang; Yang Zhang; Yanjin Zhang; Yingmei Zhang; Yunjiao Zhang; Mei Zhao; Wei-Li Zhao; Xiaonan Zhao; Yan G Zhao; Ying Zhao; Yongchao Zhao; Yu-Xia Zhao; Zhendong Zhao; Zhizhuang J Zhao; Dexian Zheng; Xi-Long Zheng; Xiaoxiang Zheng; Boris Zhivotovsky; Qing Zhong; Guang-Zhou Zhou; Guofei Zhou; Huiping Zhou; Shu-Feng Zhou; Xu-Jie Zhou; Hongxin Zhu; Hua Zhu; Wei-Guo Zhu; Wenhua Zhu; Xiao-Feng Zhu; Yuhua Zhu; Shi-Mei Zhuang; Xiaohong Zhuang; Elio Ziparo; Christos E Zois; Teresa Zoladek; Wei-Xing Zong; Antonio Zorzano; Susu M Zughaier Journal: Autophagy Date: 2016 Impact factor: 16.016
Authors: Ewelina K Wardzinski; Kamila Jauch-Chara; Sarah Haars; Uwe H Melchert; Harald G Scholand-Engler; Kerstin M Oltmanns Journal: Nutrients Date: 2022-01-14 Impact factor: 5.717
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