Divergent mutational processes distinguish hypoxic and normoxic tumours

Bhandari, Vinayak; Li, Constance H.; Bristow, Robert G.; Boutros, Paul C.

doi:10.1038/s41467-019-14052-x

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Published: 05 February 2020

Divergent mutational processes distinguish hypoxic and normoxic tumours

Nature Communications volume 11, Article number: 737 (2020) Cite this article

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An Author Correction to this article was published on 08 December 2022

This article has been updated

Abstract

Many primary tumours have low levels of molecular oxygen (hypoxia), and hypoxic tumours respond poorly to therapy. Pan-cancer molecular hallmarks of tumour hypoxia remain poorly understood, with limited comprehension of its associations with specific mutational processes, non-coding driver genes and evolutionary features. Here, as part of the ICGC/TCGA Pan-Cancer Analysis of Whole Genomes (PCAWG) Consortium, which aggregated whole genome sequencing data from 2658 cancers across 38 tumour types, we quantify hypoxia in 1188 tumours spanning 27 cancer types. Elevated hypoxia associates with increased mutational load across cancer types, irrespective of underlying mutational class. The proportion of mutations attributed to several mutational signatures of unknown aetiology directly associates with the level of hypoxia, suggesting underlying mutational processes for these signatures. At the gene level, driver mutations in TP53, MYC and PTEN are enriched in hypoxic tumours, and mutations in PTEN interact with hypoxia to direct tumour evolutionary trajectories. Overall, hypoxia plays a critical role in shaping the genomic and evolutionary landscapes of cancer.

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Introduction

Approximately half of all solid tumours are characterized by low levels of molecular oxygen (hypoxia)^1,2,3,4. Sub-regions of hypoxia can result from disrupted oxygen supply: irregular and disorganized tumour vasculature can reduce oxygen availability⁵. Hypoxia can also be caused by changes in oxygen demand: altered tumour metabolism^6,7 can increase intra-cellular demand for oxygen, potentially extending hypoxia signalling to liquid tumours. The adaptation of tumour cells to this imbalance in oxygen supply and demand is associated with poor clinical prognosis in several cancer types, attributed at least in part to hypoxia-associated genomic instability and clonal selection^{8,9,10,11,12,13,14,15,16}.

Previous work has provided insight into the molecular origins and consequences of tumour hypoxia and genomic instability. Dynamic cycling of hypoxia can select for cells with TP53 mutations and for those that are apoptosis-deficient^17,18. Indeed mutations in TP53 occur at a higher frequency in hypoxic primary tumours of at least 9 types¹⁶. The abundance of proteins involved in homologous recombination (e.g. RAD51) and non-homologous end joining (e.g. Ku70) are reduced under hypoxia, and these changes can persist for 2 days after reoxygenation^19,20,21. Genes central to efficient mismatch repair (e.g. MLH1 and MSH2) are also downregulated under hypoxia^22,23. Further, co-presence of tumour hypoxia and high genomic instability^14,15, specific cellular morphologies like intraductal and cribriform carcinoma²⁴ or specific mutations like loss of PTEN¹⁶, synergistically predict for rapid relapse after definitive local therapy in some tumour types, particularly prostate cancer. These data underscore the relationship between hypoxia and DNA repair defects, and suggest the tumour microenvironment applies a selective pressure leading to the development of specific genomic profiles.

We previously evaluated the exomic and copy-number alteration (CNA) consequences of tumour hypoxia across 19 cancer types¹⁶. However, the influence of tumour hypoxia on pan-cancer driver alterations, mutational signatures, and subclonal architectures remains unclear. To fill this gap, we calculated tumour hypoxia scores for 1188 tumours with whole-genome sequencing (WGS) and RNA sequencing, spanning 27 cancer types. Genome sequencing data was aggregated by the Pan-Cancer Analysis of Whole Genomes (PCAWG) consortium and generated by the ICGC and TCGA projects. These sequencing data were re-analyzed with standardized, high-accuracy pipelines to align to the human genome (reference build hs37d5) and identify germline variants and somatic mutations, as described previously²⁵. This sequencing data together with our high-quality hypoxia quantitation represents a powerful hypothesis-generating mechanism to suggest useful back-translational in vitro experiments and better define the hypoxia-associated mutator phenotype across cancers. We associated hypoxia with key driver alterations in coding and non-coding regions of the genome, and find hypoxia is associated with specific mutational signatures of unknown aetiology. We illustrate the joint impact of PTEN and the tumour microenvironment in influencing the evolutionary trajectory of tumours. Overall, these data highlight the genomic changes through which hypoxia drives aggressive cancers.

Results

The pan-cancer landscape of tumour hypoxia

We compiled a cohort of 1188 tumours from 27 cancer types via the PCAWG Consortium. All samples had matched tumour and reference normal WGS and tumour RNA sequencing data generated by the ICGC and TCGA projects. WGS²⁵ and RNA-sequencing²⁶ analyses were systematically carried out by centralized teams with consistent and high-accuracy bioinformatics pipelines. Normal reference samples had a mean WGS coverage of 39 reads per base-pair while coverage for tumour samples had a bimodal distribution with modes at 38 and 60 reads per base-pair²⁵. All samples underwent an extensive and systematic quality assurance process²⁵.

We used linear mixed-effect models to associate hypoxia with features of interest across cancers while adjusting for tumour purity, age, and sex^27,28. Cancer type was further incorporated as a random effect in every model, allowing us to consider a different baseline value for the feature of interest for each cancer type. As a measure of effect size we report conditional R² values, denoted as $R_{{{\mathrm{{{LMEM - C}}}}}}^2$, which reflect the variance explained by the fixed and random factors in each model²⁹. We also report marginal R² values, denoted as $R_{{{\mathrm{{{LMEM - M}}}}}}^2$, which reflect the variance explained only by the fixed factors²⁹.

We scored tumour hypoxia in all 1188 tumours using a trio of mRNA-based hypoxia signatures from Buffa³⁰, Winter³¹ and Ragnum³² (Fig. 1a, Supplementary Fig. 1a, b, Supplementary Table 1, Supplementary Data 1). Hypoxia scores from each of these independent signatures were strongly correlated (ρ = 0.71–0.88, all p < 2.2 × 10⁻¹⁶, AS89; Supplementary Fig. 1c) and consistently predicted squamous tumours of the lung (Lung-SCC), cervix (Cervix-SCC), and head (Head-SCC) as the most hypoxic (Supplementary Fig. 1d, e). Comparatively, chronic lymphocytic leukaemias (Lymph-CLL) and thyroid adenocarcinomas (Thy-AdenoCA) were the least hypoxic, consistent with previous¹⁶ reports (ρ = 0.94, p < 2.2 × 10⁻¹⁶, AS89; Fig. 1b, Supplementary Fig. 1f, h). Remarkably, subsets of patients from 23/27 cancer types have tumours with elevated hypoxia (hypoxia score > 0) and tumours consistently have elevated hypoxia compared to normal tissues (Supplementary Fig. 2a–c).

**Fig. 1: The pan-cancer landscape of tumour hypoxia.**

Considering the strong agreement between the Winter, Buffa and Ragnum hypoxia signatures (Fig. 1a, Supplementary Fig. 1c, d), we used the Buffa signature for subsequent analyses. The Buffa signature has been previously used for pan-cancer analyses and shows results consistent with those from other signatures¹⁶. We first assessed the degree of inter-tumoural heterogeneity in hypoxia that lies within individual cancer types rather than between them. Over 42% of the variance in hypoxia scores occurs within individual cancer types, highlighting the microenvironmental diversity between tumours arising in a single tissue. This variability in hypoxia score within cancer types was especially elevated in some tumour types, particularly biliary adenocarcinomas (interquartile range, IQR = 43.0; Biliary-AdenoCA), mature B-cell lymphomas (IQR = 36.0; Lymph-BNHL), lung adenocarcinomas (IQR = 34.0; Lung-AdenoCA) and breast adenocarcinomas (IQR = 32.0; Breast-AdenoCA). This was in contrast to chronic lymphocytic leukaemias (IQR = 2.0; Lymph-CLL) and prostate adenocarcinomas (IQR = 6.0; Prost-AdenoCA) where little inter-tumoural variability in hypoxia was observed. The variability in hypoxia score was not significantly associated with the median hypoxia score within cancer types (ρ = 0.20, p = 0.30, AS89; Supplementary Fig. 2d) or with sample size (ρ = 0.22, p = 0.27, AS89; Supplementary Fig. 2e). Overall, extensive heterogeneity exists in hypoxia levels within and across cancer types.

The genomic correlates of tumour hypoxia

To determine whether genomic instability arising from specific mutational classes is associated with hypoxia, we looked to identify hypoxia-associated pan-cancer mutational density and summary features³³. As a positive control, we first considered the percentage of the genome with a copy-number aberration (PGA), an engineered feature that is a surrogate for genomic instability and is associated with hypoxia across several tumour types¹⁶ (Supplementary Fig. 2f). Indeed, in this diverse pan-cancer cohort, hypoxic tumours have elevated genomic instability while controlling for cancer type, tumour purity, age and sex²⁷ (p = 2.41 × 10⁻⁸, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.022, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.57, linear mixed-effect model; Fig. 2a).

We then considered the association of hypoxia scores with 14 other metrics of the mutation density of CNAs, structural variants (SVs) and single nucleotide variants (SNVs) using linear mixed-effect models (Fig. 2a, Supplementary Fig. 2f, Supplementary Tables 2, 3). The strongest single correlate of tumour hypoxia was the total number of deletions, where patients with elevated hypoxia had more deletions (p = 1.11 × 10⁻¹⁰, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.023, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.59, linear mixed-effect model). Elevated numbers of other SVs such as duplications (p = 2.94 × 10⁻⁴, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0084, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model) and truncations (p = 3.29 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0062, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model) were also associated with high hypoxia, and we confirmed this within individual cancer types (Supplementary Fig. 3a). Other features associated with elevated hypoxia include smaller CNAs (p = 3.51 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0065, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.59, linear mixed-effect model) and more SNVs/Mbp (p = 5.55 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0054, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model). Since mutational density features can be correlated, we wanted to further test if SNVs per megabase were independently associated with hypoxia after adjusting for the total number of deletions. We created a linear mixed-effect model associating hypoxia with the number of SNVs per megabase while adjusting for cancer type, age, sex, tumour purity and the number of deletions. We also created a second model which lacked our feature of interest, SNVs per megabase, and compared the two models using an ANOVA (see the “Methods” section). The p-value for this comparison was 0.011, suggesting that the number of SNVs per megabase are associated with hypoxia independent of the number of deletions (and other potential confounders included in the models). Overall, hypoxia is associated with increased numbers of most types of somatic mutations.

Considering the strong association of hypoxia with mutational density, we next looked to determine if these were only general effects or selectively affected specific genes or chromosome regions. We leveraged a catalogue of 653 driver mutations²⁵, with CNA, SV and SNV data available for 1096 patients. In cases where a patient had multiple mutations in the same gene (e.g. a CNA and an SNV) we denoted these as compound events. We again used linear-mixed effect models to associate hypoxia with each driver feature across cancers (Fig. 2b). Adjusting for cancer type, tumour purity, age and sex, 10 driver events were associated with hypoxia across cancers (FDR < 0.10, linear-mixed effect models; Supplementary Fig. 2f, Supplementary Table 4). Tumours with mutations in BCL2 (FDR = 7.56 × 10⁻¹⁵, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.045, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.62, linear-mixed effect model) showed lower levels of hypoxia compared to those without. All alterations of BCL2 in this cohort were SVs, so it is important to note that this association could not be identified from previous exome-sequencing data. Similarly, mutations in the tumour suppressor TP53 were associated with elevated hypoxia across cancers (FDR = 1.97 × 10⁻¹², $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.043, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.59, linear-mixed effect model), consistent with previous descriptions of hypoxia-mediated selection of TP53-mutated cells¹⁷ and elevated hypoxia in breast cancers with TP53 mutations¹⁶. We also confirmed this association within individual cancer types (Supplementary Fig. 3b). Mutations of the oncogene MYC (FDR = 1.07 × 10⁻⁴, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.016, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear-mixed effect model) and tumour suppressor PTEN (FDR = 1.50 × 10⁻², $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0098, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.59, linear mixed-effect model) were also associated with elevated hypoxia. Alterations in mitochondrial genes³⁴ were not significantly associated with tumour hypoxia (Supplementary Fig. 3c). Thus, hypoxia is associated with both broad elevation of mutation density of most types of somatic variation, along with a consistent signature of alterations in oncogenes and tumour suppressors across cancers.

Hypoxia-associated mutational signatures

Previous work has used nonnegative matrix factorization to identify distinct mutational processes in cancer cells from endogenous and exogenous agents³⁵. To identify hypoxia-associated mutational processes, we tested if hypoxia score was associated with the proportion of mutations attributed to each mutational signature using linear-mixed effect models. Of the 65 single base substitution (SBS) signatures tested, nine showed differential activity in hypoxic tumours compared to non-hypoxic ones, while controlling for cancer type, tumour purity, age and sex (FDR < 0.10, linear mixed-effect models; Fig. 3a, Supplementary Table 5). Of these, six were more active and three less active in tumours with elevated hypoxia. Since previous work has shown that DNA repair is impaired under hypoxia, it was not surprising to observe that a higher proportion of mutations were attributed to SBS3 (related to defective homologous recombination-based repair) in tumours with elevated hypoxia score (FDR = 1.98 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.016, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear-mixed effect model). Further, SBS6 (FDR = 1.98 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0086, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.61, linear-mixed effect model) and SBS21 (FDR = 4.31 × 10⁻², $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0051, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.61, linear-mixed effect model), both related to defective DNA mismatch repair, had a higher proportion of attributed mutations with increasing hypoxia. A lower proportion of mutations were also attributed to SBS1, previously related to the deamination of 5-methylcytosine, with increasing hypoxia (FDR = 8.33 × 10⁻⁸, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.033, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.61, linear-mixed effect model).

Intriguingly, hypoxia was also associated with a number of SBS signatures with unknown aetiology (Fig. 3b). The strongest of these was SBS5, where elevated hypoxia was associated with a significantly lower proportion of mutations attributed to the signature (FDR = 1.43 × 10⁻⁶, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.022, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.59, linear-mixed effect model). A significantly lower proportion of mutations were also attributed to SBS12 with increasing hypoxia score (FDR = 4.31 × 10⁻², $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0066, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear-mixed effect model). In contrast, a higher proportion of mutations were attributed to SBS17a (FDR = 4.80 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0072, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.61, linear mixed-effect model) and SBS17b (FDR = 2.83 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.079, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.61, linear mixed-effect model) with increasing hypoxia.

Analysis of small insertion and deletion (ID) signatures illustrated a similar story. Of the 17 ID signatures analyzed, the activity of 5 were associated with tumour hypoxia scores while controlling for cancer type, tumour purity, age and sex (FDR < 0.10, linear mixed-effect models; Fig. 3b, Supplementary Table 6). Of these, 3 were more active in tumours with elevated hypoxia while 2 were less active in them. The defective homologous recombination signature ID6 (FDR = 5.76 × 10⁻⁵, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.015, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model) and defective DNA mismatch repair signature ID2 (FDR = 7.06 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.011, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.61, linear mixed-effect model) had a higher proportion of attributed mutations as hypoxia score increased. Several signatures with unknown aetiology were also significantly associated with hypoxia score, including ID5 (FDR = 1.54 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.016, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model) and ID9 (FDR = 7.06 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0068, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model). These data suggest that oxygen levels play a direct or indirect role in the accumulation of specific mutations in cancer cells that are reflected by these signatures.

The subclonal hallmarks of tumour hypoxia

State-of-the-art methods for subclonal reconstruction rely on WGS data³⁶, making the PCAWG dataset ideal for understanding the evolutionary pressures imposed by hypoxia. We and others have shown that some mutations consistently occur early during tumourigenesis while others occur later and that hypoxia is associated with CNAs occurring early in localized prostate cancer^16,37,38. To explore if this interaction between the tumour microenvironment and mutational landscape exists more broadly in cancer, we assessed if hypoxia was related to the number of clonal or subclonal mutations across 1188 tumours from 27 cancer types³⁸. Clonal mutations are common to all cells in a tumour while subclonal ones are only present in a subpopulation of cells. We found that elevated hypoxia was significantly associated with an increased number of clonal alterations across cancers (Bonferroni-adjusted p = 4.65 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0074, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model; Fig. 4a, Supplementary Table 7), particularly clonal SVs (p = 1.17 × 10⁻⁵, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.013, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model). In contrast, tumour hypoxia was not significantly associated with the number of subclonal alterations (Bonferroni-adjusted p = 0.28, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0039, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model; Fig. 4a). Further, consistent with findings in prostate cancer¹⁶, hypoxia was not associated with the number of subclones detected (Bonferroni-adjusted p = 0.14, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.0051, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.60, linear mixed-effect model; Fig. 4a). These data suggest that hypoxia applies a selective pressure on tumours during their early evolution, prior to subclonal diversification.

Next, we assessed if the mutational background of a tumour together with its oxygenation level was linked to its evolutionary trajectory. We previously demonstrated that patients with hypoxic polyclonal prostate tumours with loss of the tumour suppressor PTEN tend to have a poor prognosis¹⁶. Indeed, here we observed a significant interaction between tumour hypoxia and loss of PTEN in predicting subclonal architecture (p_interaction = 8.39 × 10⁻³, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.60, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.87, linear mixed-effect model; Fig. 4b). Specifically, tumours with both of these features tend to have a polyclonal architecture across cancers. The downstream impact of this interaction between the genome and the tumour microenvironment was observed in RNA data: tumours with both altered PTEN and elevated hypoxia had the lowest abundance of PTEN mRNA (p = 4.63 × 10⁻¹⁴, $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ = 0.054, $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ = 0.47, linear mixed-effect model; Fig. 4c). Thus, the evolutionary trajectory of a tumour may be driven by the presence of a mutation in a specific microenvironmental context (Fig. 4d).

Discussion

Hypoxia is a feature of many solid and liquid tumours and is associated with aggressive disease. We calculated hypoxia scores for 1188 tumours from 27 cancer types and showed the vast heterogeneity that exists in this microenvironmental feature within and across cancer types. This reinforces previous pushes for careful patient selection in prospective trials of hypoxia-targeting agents¹⁶.

For the first time, we characterized the pan-cancer whole-genome correlates of tumour hypoxia. We show the broad influence of the hypoxia-associated mutator phenotype: elevated hypoxia is associated with increased mutational load across all mutational classes (i.e. CNAs, SVs and SNVs). This supports previous in vitro work that demonstrated the contextual synthetic lethality of PARP inhibition in cells with defective DNA repair due to hypoxia³⁹. Regarding this co-occurrence of genomic instability and hypoxia, our group¹⁶ and others⁴⁰ have previously described this metabolic reprogramming as a series of distinct genomic alterations. This is supported by our finding that alterations in TP53, MYC and PTEN are more common in tumours with elevated hypoxia across cancers. Supporting these findings, previous in vitro work has shown that heterogeneous populations of cells where a small subpopulation have mutant TP53 can rapidly expand under cycling hypoxia to become the major subpopulation due to deficient apoptosis and selection¹⁷. We have also previously shown that tumours with TP53 mutations have elevated hypoxia within individual breast cancer subtypes, confirming that this association is not simply reflecting previously described molecular subtypes¹⁶.

Our study cannot conclusively say whether hypoxia exerts a selective pressure that enriches for specific genomic alterations or if these genomic changes directly result in hypoxia. Experimental studies of single genes support that both effects may contribute to the associations we describe^{17,22,41,42,43}. While we have not specifically included in vitro experimental validation data in this report, we and others have previously validated associations first revealed by analysis of mRNA-based hypoxia signatures. For example, our group previously described microRNA-133a-3p as a hypoxia-associated miRNA in prostate cancer based on mRNA signature-based associations across multiple independent datasets¹⁶. We went on to validate that microRNA-133a-3p was indeed induced under hypoxia in multiple prostate cancer cell lines and confirmed its capacity to modulate cell proliferation and invasion. Similarly, Ye et al. applied hypoxia signatures to 10 independent datasets of cell lines and primary tumour fragments under hypoxia and normoxia⁴⁴. These 10 datasets represented seven cancer types and within each dataset samples under hypoxia showed higher hypoxia scores compared to normoxic samples. Further, they generated predictions of drugs that would be more or less potent under hypoxia and validated four drug–hypoxia interactions in vitro. These data illustrate that hypoxia signatures applied to large cohorts of primary tumours can generate reliable hypotheses, many of which have been validated in controlled systems. However, some aerobic cancer cells may also mimic the biological state of hypoxia (i.e., pseudohypoxia) and this may affect signature-derived hypoxia estimates. Further, while pimonidazole (which was used to develop the Ragnum hypoxia signature³²) reflects oxygen tensions below 10 mmHg (1.3% O₂), it is difficult to directly relate hypoxia signature scores with oxygen tension⁴⁵. Overall, hypoxia signalling can be distinct from microenvironmental hypoxia and this remains a critical caveat of this study.

Diving into the mutational processes related to hypoxia, we confirmed that several SBS and small indel signatures related to impaired DNA repair were associated with hypoxia. This raises the potential confounder that because hypoxic tumours have more mutations, we have more power to detect related mutational signatures. However, we demonstrated that hypoxia is indeed strongly associated with many mutational signatures with unknown aetiology, particularly SBS5, which is found in nearly all cancer types. Modelling these associations in vitro is particularly difficult and these data provide a high confidence measure of the mutational signatures that may be directly or indirectly driven by tumour oxygen levels. It is difficult to disentangle the timing of these events: whether a specific driver mutation gives rise to a specific mutational signature or if these are separate processes. Better mapping of the evolutionary timing of hypoxia will be particularly important in addressing this question and the advent of hypoxia signatures may facilitate future studies in this area.

We observed a significant association between elevated hypoxia and the number of clonal mutations. This supports the idea that hypoxia is an early event in cancer, as we have suggested previously¹⁶, and other models that link hypoxia to genomic instability and downstream clonal selection^20,42. Previous work has also demonstrated that patients with allelic loss of PTEN and elevated hypoxia rapidly relapse after definitive treatment for localized prostate cancer¹⁶. Here, we showed that tumours with alterations in PTEN and elevated hypoxia are enriched for a polyclonal tumour architecture. This illustrates the joint influence of the tumour mutational landscape and microenvironment in guiding evolutionary trajectories across cancers. Further, these data suggest that increased subclonal diversification may be a novel route via which PTEN drives aggressive tumour phenotypes, in concert with tumour hypoxia, and this can be better defined with future back-translational in vitro experiments. The PCAWG dataset is the largest publicly available pan-cancer dataset to date and this limits our ability to validate our discoveries in independent datasets. Ultimately, it will be necessary to validate our findings in large, independent cohorts. Hypotheses generated here, particularly those around hypoxia and tumour evolution, will require long term, systematic in vitro modelling and will be the subject of future studies. Overall, this work shows that a hypoxic tumour microenvironment is associated with specific mutational processes and distinct somatic mutational profiles, and may direct the subclonal architecture of cancers.

Methods

Pan-cancer hypoxia scoring

Hypoxia scores were calculated for all 1188 tumours with mRNA abundance data (FPKM with upper-quartile normalization) using mRNA-abundance-based signatures of tumour hypoxia developed previously by Winter et al.³¹, Buffa et al. ³⁰ and Ragnum et al. ³², as described previously^14,16 (Supplementary Data 1). Briefly, patients with the top 50% of mRNA abundance values for each gene in a signature were given a score of +1. Patients with the bottom 50% of mRNA abundance values for that gene were given a score of −1. This was repeated for every gene in the signature to generate a hypoxia score for each patient, and this process was repeated for each of the three signatures used in the study. High scores suggest that the tumour was hypoxic and low scores are indicative of normoxia.

Hypoxia score comparison

To compare hypoxia scores generated by the different signatures, the median hypoxia score was calculated for each of the PCAWG cancer types based on each signature. The median hypoxia scores from each signature were then scaled from +1 to −1 using the plotrix package (v3.7). Scaled median hypoxia values for the PCAWG cancer types were also compared to scaled median hypoxia values from previously published¹⁶ TCGA data between comparable cancer groups. The groups compared are as follows (PCAWG cancer type and TCGA cancer type): Bladder-TCC and BLCA; Breast-AdenoCA and BRCA; Cervix-SCC and CESC; CNS-GBM and GBM; ColoRect-AdenoCA and COADREAD; Head-SCC and HNSC; Kidney-RCC and KIRC; Liver-HCC and LIHC; Lung-AdenoCA and LUAD; Lung-SCC and LUSC; Ovary-AdenoCA and OV; Panc-AdenoCA and PAAD; Prost-AdenoCA and PRAD; Skin-Melanoma and SKCM; Thy-AdenoCA and THCA; Uterus-Adeno and UCEC. Of the 27 cancer types in PCAWG, 11 (Cervix-AdenoCA, Stomach-AdenoCA, Eso-AdenoCA, Breast-LobularCA, SoftTissue-Leiomyo, Lymph-BNHL, SoftTissue-Liposarc, Biliary-AdenoCA, Kidney-ChRCC, CNS-Oligo and Lymph-CLL) did not have hypoxia data from comparable cancers in TCGA and were not used for the comparison¹⁶. For Spearman’s correlations, p-values were calculated using algorithm AS89.

Tumour vs. normal hypoxia comparison

Previously calculated pan-cancer tumour hypoxia scores were gathered for 7791 independent tumours from 19 cancer types based on the Buffa hypoxia signature¹⁶. Hypoxia scores were then calculated for all normal tissue samples related to the 19 cancer types with hypoxia scores (n = 640 independent normal tissue samples). Tumour hypoxia scores were compared to normal tissue hypoxia scores for each tissue type where at least 15 normal tissue samples were available (Mann–Whitney U-test). A total of 5649 independent tumours and 625 independent normal tissue samples were evaluated in the comparisons.

Linear mixed-effect models

We used linear mixed-effect models to associate hypoxia with features of interest (e.g., PGA, TP53 mutational status, etc.) across cancers using the lme4 package (v1.1-17). For each feature of interest, we compared a full model (i.e., a model with the feature of interest) to a null model (i.e. a model without the feature of interest) using an ANOVA to determine if hypoxia was significantly associated with the feature of interest across cancers. A generic example of this is shown below with Eqs. (1) and (2):

$${{{\mathrm{{full}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{feature}}}}} + {{{\mathrm{{purity}}}}} + {{{\mathrm{{age}}}}} + {{{\mathrm{{sex}}}}} + \left( {1\left| {{{{\mathrm{{cancer}}}}}} \right.} \right)$$

(1)

$${{{\mathrm{{null}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{purity}}}}} + {{{\mathrm{{age}}}}} + {{{\mathrm{{sex}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(2)

All models were adjusted for tumour purity, patient age and sex^27,28. Cancer type was incorporated as a random effect in every model. This allowed us to consider a different baseline value for the feature of interest for each cancer type. For each model a conditional R² value is reported ($R_{{{\mathrm{{{LMEM - C}}}}}}^2$) which reflects the variance explained by the fixed and random factors²⁹. We also report marginal R² values for each model ($R_{{{\mathrm{{{LMEM - M}}}}}}^2$) which reflect the variance explained by the fixed factors only²⁹. $R_{{{\mathrm{{{LMEM - C}}}}}}^2$ and $R_{{{\mathrm{{{LMEM - M}}}}}}^2$ values were calculated as described previously²⁹.

All model diagnostics were done using the DHARMa package (0.2.0) which uses a simulation-based approach to create standardized residuals⁴⁶. For each model, scaled residuals were generated using the simulateResiduals function. The full model was used as the input for fittedModel parameter and 1000 simulations were run. For correctly specified models, the scaled residuals were expected to be uniformly distributed and this was verified for each full model. We also compared the standardized residuals to the rank transformed predicted values to assess deviations from uniformity for each full model.

Mutational density analysis

Previously published data for 15 mutational density and summary features were downloaded for 1188 tumours³³. We used linear mixed-effect models to associate each feature with hypoxia score across cancers and compared each full model with a null model. Cancer type was incorporated as a random effect in each model while tumour purity, age and sex were incorporated as fixed effects. Tumours belonging to cancer types with fewer than 15 samples were excluded from the analysis. A Bonferroni p-value adjustment was applied to the p-values from linear mixed-effect modelling since fewer than 20 tests were conducted. All models were adjusted for tumour purity based on previously published purity data³³. The full model for evaluating PGA is shown below as an example as follows:

$${{{\mathrm{{full}}}}}_{{{\mathrm{{{PGA}}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{PGA + purity + age + sex}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(3)

To assess if SNVs per megabase were independently associated with hypoxia after adjusting for the total number of deletions we created two linear mixed-effect models. The full model associated hypoxia with SNVs per megabase while adjusting for cancer type, age, sex, tumour purity and the number of deletions (Eq. (4)). For comparison, a null model was created without our feature of interest, SNVs per megabase (Eq. (5)). The two models were compared using an ANOVA.

$$\begin{array}{l}{{{\mathrm{{full}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{SNVs}}}}}\,{{{\mathrm{{per}}}}}\,{{{\mathrm{{megabase + purity + age + sex}}}}}\\ + \, {{{\mathrm{{total}}}}}\,{{{\mathrm{{deletions}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)\end{array}$$

(4)

$${{{\mathrm{{null}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{purity + age + sex +}}}}} {{{\mathrm{{total}}}}}\, {{{\mathrm{{deletions}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(5)

Driver mutations analysis

Data for driver mutations was first summarized at the gene level for 1096 tumours with previously published driver mutation data²⁵. For each of the 653 driver features, we summarized if a patient had an SNV, CNA or SV. Some tumours had more than one type of event in a gene (e.g. a CNA and an SNV) and these events were classified as compound events. We then used linear mixed-effect models to associate the mutational status of each gene with hypoxia score and compared each full model with a null model. Cancer type was incorporated as a random effect in each model while tumour purity, age and sex were incorporated as fixed effects. The driver mutation analysis did not specifically consider the type of mutation in the gene and only considered if the gene had a mutation or was wildtype. Tumours belonging to cancer types with fewer than 15 samples were excluded from the analysis. An FDR adjustment was applied to the p-values from linear mixed-effect modelling. The full model for evaluating PTEN is shown below as an example as follows:

$${{{\mathrm{{full}}}}}_{{{\mathrm{{{PTEN}}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{PTEN + purity + age + sex}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(6)

Mutational signature analysis

Previously published data for mutations attributed to various specific signatures was downloaded for 1188 tumours³⁵. For each tumour, we calculated the proportion of total mutations attributed to each mutational signature. The proportion of mutations attributed to each signature were calculated by dividing the number of mutations attributed to each signature by the total number of mutations in the tumour. We used linear mixed-effect models to associate the proportion of mutations attributed to each signature with hypoxia score and compared each full model with a null model. Cancer type was incorporated as a random effect in each model while purity, age and sex were incorporated as fixed effects. Tumours belonging to cancer types with fewer than 15 samples were excluded from the analysis. An FDR adjustment was applied to the p-values from linear mixed-effect modelling. The full model for SBS1 is shown below as an example as follows:

$${{{\mathrm{{full}}}}}_{{{\mathrm{{{SBS1}}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{SBS1 + purity + age + sex}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(7)

Subclonality analysis

Previously reported³⁸ subclonal reconstruction data was used to summarize the number of clonal and subclonal mutations in all 1188 tumours. We used linear mixed-effect models to associate the number of these timed mutations with hypoxia score and compared each full model with a null model. Cancer type was incorporated as a random effect in each model while purity, age and sex were incorporated as fixed effects. Tumours belonging to cancer types with fewer than 15 samples were excluded from the analysis. A Bonferroni adjustment was applied to the p-values from linear mixed-effect modelling since fewer than 20 tests were conducted.

The number of subclones was calculated for all 1188 tumours based on the number of clusters of cells identified in each sample. A linear mixed-effects model was used to associate the number of subclones with hypoxia score and this model was compared to a null model. Cancer type was incorporated as a random effect while purity, age and sex were incorporated as fixed effects. Tumours belonging to cancer types with fewer than 15 samples were excluded from the analysis. The full model for associating the number of subclones with hypoxia score is shown below as follows:

$${{{\mathrm{{full}}}}}_{{{\mathrm{{{subclones}}}}}} = {{{\mathrm{{hypoxia}}}}}\sim {{{\mathrm{{subclones + purity + age + sex }}}}}+ \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(8)

Patients with only one identified cluster of cells were defined as monoclonal and patients with more than one identified cluster of cells were defined as polyclonal³⁷. Hypoxia scores were median dichotomized to classify patients as hypoxic or normoxic. To test for an interaction between tumour hypoxia and PTEN mutational status in selecting for a particular subclonal architecture, we used linear mixed-effect models together with an ANOVA. An interaction model was first created where the relationship between the hypoxia scores and PTEN mutational status was modelled as an interaction (Eq. (9)). An additive model was also created where the relationship between hypoxia scores and PTEN mutational status was modelled in an additive manner (Eq. (10)):

$${{{\mathrm{{interaction}}}}} = {{{\mathrm{{clonality}}}}}\sim {{{\mathrm{{hypoxia}}}}} \ast {{{\mathrm{{PTEN + purity + age + sex}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(9)

$${{{\mathrm{{additive}}}}} = {{{\mathrm{{clonality}}}}}\sim {{{\mathrm{{hypoxia + PTEN + purity + age + sex}}}}} + \left( {1|{{{\mathrm{{cancer}}}}}} \right)$$

(10)

The two models were compared using an ANOVA to test if hypoxia scores significantly interact with PTEN mutational status. Tumours belonging to cancer types with fewer than 15 samples were excluded from the analysis. The full model diagnostics were carried out using the DHARMa package, as described above.

All data analysis was performed in the R statistical environment (v3.4.3). Data visualization was performed using the BPG package⁴⁷ (v5.9.1). Figures were compiled using Inkscape (v0.91).

Reporting summary

Further information on research design is available in the Nature Research Reporting Summary linked to this article.

Data availability

Aligned sequencing data, somatic and germline variant calls, single nucleotide variants, indels, copy-number alterations, structural variants, mutational signatures, subclonal reconstructions, transcript abundances and other core data generated by the PCAWG Consortium are described here²⁵ and available for download at dcc.icgc.org/releases/PCAWG. Additional information on accessing the data, including raw read files, can be found at docs.icgc.org/pcawg/data. In accordance with the data access policies of the ICGC and TCGA projects, most molecular, clinical and specimen data are in an open tier which does not require approval for access.

To access potentially identifiable information, such as germline alleles and underlying sequencing data, researchers will need to apply to the TCGA Data Access Committee (DAC) via dbGaP (dbgap.ncbi.nlm.nih.gov/aa/wga.cgi?page = login) for access to the TCGA portion of the dataset, and to the ICGC Data Access Compliance Office (DACO; icgc.org/daco) for the ICGC portion of the dataset. To access somatic single nucleotide variant data derived from TCGA donors, researchers will also need to obtain dbGaP authorization.

The analyses in this paper used several PCAWG datasets that were derived from the raw sequencing data and variant calls. The individual datasets are available at Synapse (synapse.org) and are denoted with accession numbers starting with syn. All of these datasets are also mirrored at dcc.icgc.org. The datasets encompass: clinical data for each patient, including information about age and sex (syn10389158, dcc.icgc.org/releases/PCAWG/clinical_and_histology); histopathology data for each tumour (syn10389164, dcc.icgc.org/releases/PCAWG/clinical_and_histology); mRNA abundance data for each tumour (syn5553991, dcc.icgc.org/releases/PCAWG/transcriptome/gene_expression); driver mutations for each tumour spanning all classes of variants (syn11639581, dcc.icgc.org/releases/PCAWG/driver_mutations); single base substitution and small indel signature exposures for each tumour (syn8366024, dcc.icgc.org/releases/PCAWG/mutational_signatures); subclonal architecture, tumour evolution and tumour purity data for each tumour (syn8272483, dcc.icgc.org/releases/PCAWG/subclonal_reconstruction).

Code availability

The core computational pipelines used by the PCAWG Consortium for alignment, quality control and variant calling are available to the public at dockstore.org/search?search = pcawg under the GNU General Public License v3.0, which allows for reuse and distribution. Code specific to this manuscript is available upon request.

Change history

08 December 2022
A Correction to this paper has been published: https://doi.org/10.1038/s41467-022-32339-4

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Acknowledgements

The authors gratefully thank Dr. Marianne Koritzinsky for insightful suggestions and thank Jenna E. van Leeuwen for visualization support. The authors also thank all members of the Boutros and Bristow labs for helpful suggestions. We acknowledge the contributions of the many clinical networks across ICGC and TCGA who provided samples and data to the PCAWG Consortium, and the contributions of the Technical Working Group and the Germline Working Group of the PCAWG Consortium for collation, realignment and harmonised variant calling of the cancer genomes used in this study. We thank the patients and their families for their participation in the individual ICGC and TCGA projects. This study was conducted with the support of Movember funds through Prostate Cancer Canada, and with the additional support of the Ontario Institute for Cancer Research, funded by the Government of Ontario. This work was supported by Prostate Cancer Canada and is proudly funded by the Movember Foundation Team Grant T2013 and #RS2014-01. Paul C. Boutros was supported by a Terry Fox Research Institute New Investigator Award, a Prostate Cancer Canada Rising Star Fellowship, a CIHR New Investigator Award, a CIHR Project Grant, the Government of Canada through Genome Canada and the Ontario Genomics Institute (OGI-125), Canadian Cancer Society (grant #705649) and by the University of California. This work has been funded by a fellowship from the Canadian Institutes of Health Research to Vinayak Bhandari. Constance Li was supported by an award from the Government of Ontario. The authors gratefully thank the Princess Margaret Cancer Centre Foundation and Radiation Medicine Programme Academic Enrichment Fund for support (to Robert G. Bristow). This work was supported by a Terry Fox Research Institute Programme Project Grant. Robert G. Bristow is a recipient of a Canadian Cancer Society Research Scientist Award. Laboratory work for R.G.B is supported by the CRUK Manchester Institute through Cancer Research UK. This work was supported by the NIH/NCI through award number P30CA016042.

Author information

Authors and Affiliations

Department of Medical Biophysics, University of Toronto, Toronto, ON, Canada
Vinayak Bhandari, Constance H. Li & Paul C. Boutros
Ontario Institute for Cancer Research, Toronto, Canada
Vinayak Bhandari & Constance H. Li
Department of Human Genetics, University of California, Los Angeles, USA
Constance H. Li & Paul C. Boutros
Division of Cancer Sciences, Faculty of Biology, Health and Medicine, University of Manchester, Manchester, UK
Robert G. Bristow
The Christie NHS Foundation Trust, Manchester, UK
Robert G. Bristow
CRUK Manchester Institute and Centre, Manchester, UK
Robert G. Bristow
Department of Pharmacology and Toxicology, University of Toronto, Toronto, Canada
Paul C. Boutros
Vector Institute for Artificial Intelligence, Toronto, Canada
Paul C. Boutros
Department of Urology, University of California, Los Angeles, USA
Paul C. Boutros
Jonsson Comprehensive Cancer Centre, University of California Los Angeles, Los Angeles, USA
Paul C. Boutros
Institute for Precision Health, University of California Los Angeles, Los Angeles, USA
Paul C. Boutros
Applied Tumor Genomics Research Program, Research Programs Unit, University of Helsinki, Helsinki, Finland
Lauri A. Aaltonen
Wellcome Sanger Institute, Wellcome Genome Campus, Hinxton, UK
Federico Abascal, David J. Adams, Ludmil B. Alexandrov, Sam Behjati, Shriram G. Bhosle, David T. Bowen, Adam P. Butler, Peter J. Campbell, Peter Clapham, Helen Davies, Kevin J. Dawson, Stefan C. Dentro, Serge Serge, Erik Garrison, Mohammed Ghori, Dominik Glodzik, Jonathan Hinton, David R. Jones, Young Seok Ju, Stian Knappskog, Barbara Kremeyer, Henry Lee-Six, Daniel A. Leongamornlert, Yilong Li, Sancha Martin, Iñigo Martincorena, Ultan McDermott, Andrew Menzies, Thomas J. Mitchell, Sandro Morganella, Jyoti Nangalia, Jonathan Nicholson, Serena Nik-Zainal, Sarah O’Meara, Elli Papaemmanuil, Keiran M. Raine, Manasa Ramakrishna, Kamna Ramakrishnan, Nicola D. Roberts, Rebecca Shepherd, Lucy Stebbings, Michael R. Stratton, Maxime Tarabichi, Jon W. Teague, Ignacio Vázquez-García, David C. Wedge, Lucy Yates, Jorge Zamora & Xueqing Zou
Memorial Sloan Kettering Cancer Center, New York, NY, USA
Adam Abeshouse, Hikmat Al-Ahmadie, Gunes Gundem, Zachary Heins, Jason Huse, Douglas A. Levine, Eric Minwei Liu & Angelica Ochoa
Genome Science Division, Research Center for Advanced Science and Technology, University of Tokyo, Tokyo, Japan
Hiroyuki Aburatani, Genta Nagae, Akihiro Suzuki, Kenji Tatsuno & Shogo Yamamoto
Department of Surgery, University of Chicago, Chicago, IL, USA
Nishant Agrawal
Department of Surgery, Division of Hepatobiliary and Pancreatic Surgery, School of Medicine, Keimyung University Dongsan Medical Center, Daegu, South Korea
Keun Soo Ahn & Koo Jeong Kang
Department of Oncology, Gil Medical Center, Gachon University, Incheon, South Korea
Sung-Min Ahn
Hiroshima University, Hiroshima, Japan
Hiroshi Aikata, Koji Arihiro, Kazuaki Chayama, Yoshiiku Kawakami & Hideki Ohdan
Department of Bioinformatics and Computational Biology, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Rehan Akbani, Shaolong Cao, Yiwen Chen, Zechen Chong, Yu Fan, Jun Li, Han Liang, Wenyi Wang, Yumeng Wang & Yuan Yuan
University of Texas MD Anderson Cancer Center, Houston, TX, USA
Kadir C. Akdemir & Ken Chen
King Faisal Specialist Hospital and Research Centre, Al Maather, Riyadh, Saudi Arabia
Sultan T. Al-Sedairy
Bioinformatics Unit, Spanish National Cancer Research Centre (CNIO), Madrid, Spain
Fatima Al-Shahrour & Elena Piñeiro-Yáñez
Bioinformatics Core Facility, University Medical Center Hamburg, Hamburg, Germany
Malik Alawi
Heinrich Pette Institute, Leibniz Institute for Experimental Virology, Hamburg, Germany
Malik Alawi & Adam Grundhoff
Ontario Tumour Bank, Ontario Institute for Cancer Research, Toronto, ON, Canada
Monique Albert & John Bartlett
Department of Pathology, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Kenneth Aldape, Russell R. Broaddus, Bogdan Czerniak, Adel El-Naggar, Savitri Krishnamurthy, Alexander J. Lazar & Xiaoping Su
Laboratory of Pathology, Center for Cancer Research, National Cancer Institute, Bethesda, MD, USA
Kenneth Aldape
Department of Cellular and Molecular Medicine and Department of Bioengineering, University of California San Diego, La Jolla, CA, USA
Ludmil B. Alexandrov & Erik N. Bergstrom
UC San Diego Moores Cancer Center, San Diego, CA, USA
Ludmil B. Alexandrov, Erik N. Bergstrom & Olivier Harismendy
Canada’s Michael Smith Genome Sciences Centre, BC Cancer, Vancouver, BC, Canada
Adrian Ally, Miruna Balasundaram, Reanne Bowlby, Denise Brooks, Rebecca Carlsen, Eric Chuah, Noreen Dhalla, Robert A. Holt, Steven J. M. Jones, Katayoon Kasaian, Darlene Lee, Haiyan Irene Li, Yussanne Ma, Marco A. Marra, Michael Mayo, Richard A. Moore, Andrew J. Mungall, Karen Mungall, A. Gordon Robertson, Sara Sadeghi, Jacqueline E. Schein, Payal Sipahimalani, Angela Tam, Nina Thiessen & Tina Wong
Sir Peter MacCallum Department of Oncology, Peter MacCallum Cancer Centre, University of Melbourne, Melbourne, VIC, Australia
Kathryn Alsop, David D. L. Bowtell, Elizabeth L. Christie, Dariush Etemadmoghadam, Sian Fereday, Dale W. Garsed, Linda Mileshkin, Chris Mitchell, Mark Shackleton, Heather Thorne & Nadia Traficante
Centre for Research in Molecular Medicine and Chronic Diseases (CiMUS), Universidade de Santiago de Compostela, Santiago de Compostela, Spain
Eva G. Alvarez, Alicia L. Bruzos, Bernardo Rodriguez-Martin, Javier Temes, Jose M. C. Tubio & Jorge Zamora
Department of Zoology, Genetics and Physical Anthropology, (CiMUS), Universidade de Santiago de Compostela, Santiago de Compostela, Spain
Eva G. Alvarez, Alicia L. Bruzos, Bernardo Rodriguez-Martin, Javier Temes, Jose M. C. Tubio & Jorge Zamora
The Biomedical Research Centre (CINBIO), Universidade de Vigo, Vigo, Spain
Eva G. Alvarez, Alicia L. Bruzos, Bernardo Rodriguez-Martin, Marta Tojo, Jose M. C. Tubio & Jorge Zamora
Royal National Orthopaedic Hospital - Bolsover, London, UK
Fernanda Amary
Department of Genomic Medicine, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Samirkumar B. Amin, P. Andrew Futreal & Alexander J. Lazar
Quantitative and Computational Biosciences Graduate Program, Baylor College of Medicine, Houston, TX, USA
Samirkumar B. Amin, Han Liang & Yumeng Wang
The Jackson Laboratory for Genomic Medicine, Farmington, CT, USA
Samirkumar B. Amin, Joshy George & Lucas Lochovsky
Genome Informatics Program, Ontario Institute for Cancer Research, Toronto, ON, Canada
Brice Aminou, Niall J. Byrne, Aurélien Chateigner, Nodirjon Fayzullaev, Vincent Ferretti, George L. Mihaiescu, Hardeep K. Nahal-Bose, Brian D. O’Connor, B. F. Francis Ouellette, Marc D. Perry, Kevin Thai, Qian Xiang, Christina K. Yung & Junjun Zhang
Institute of Human Genetics, Christian-Albrechts-University, Kiel, Germany
Ole Ammerpohl, Andrea Haake, Cristina López, Julia Richter & Rabea Wagener
Institute of Human Genetics, Ulm University and Ulm University Medical Center, Ulm, Germany
Ole Ammerpohl, Sietse Aukema, Cristina López, Reiner Siebert & Rabea Wagener
Queensland Centre for Medical Genomics, Institute for Molecular Bioscience, University of Queensland, St. Lucia, Brisbane, QLD, Australia
Matthew J. Anderson, Timothy J. C. Bruxner, Angelika N. Christ, J. Lynn Fink, Ivon Harliwong, Karin S. Kassahn, David K. Miller, Alan J. Robertson & Darrin F. Taylor
Salford Royal NHS Foundation Trust, Salford, UK
Yeng Ang, Hsiao-Wei Chen, Ritika Kundra & Francisco Sanchez-Vega
Department of Surgery, Pancreas Institute, University and Hospital Trust of Verona, Verona, Italy
Davide Antonello, Claudio Bassi, Narong Khuntikeo, Luca Landoni, Giuseppe Malleo, Giovanni Marchegiani, Neil D. Merrett, Marco Miotto, Salvatore Paiella, Antonio Pea, Paolo Pederzoli, Roberto Salvia, Jaswinder S. Samra, Elisabetta Sereni & Samuel Singer
Molecular and Medical Genetics, OHSU Knight Cancer Institute, Oregon Health and Science University, Portland, OR, USA
Pavana Anur, Myron Peto & Paul T. Spellman
Department of Molecular Oncology, BC Cancer Research Centre, Vancouver, BC, Canada
Samuel Aparicio
The McDonnell Genome Institute at Washington University, St. Louis, MO, USA
Elizabeth L. Appelbaum, Matthew H. Bailey, Matthew G. Cordes, Li Ding, Catrina C. Fronick, Lucinda A. Fulton, Robert S. Fulton, Kuan-lin Huang, Reyka Jayasinghe, Elaine R. Mardis, R. Jay Mashl, Michael D. McLellan, Christopher A. Miller, Heather K. Schmidt, Jiayin Wang, Michael C. Wendl, Richard K. Wilson & Tina Wong
University College London, London, UK
Elizabeth L. Appelbaum, Jonathan D. Kay, Helena Kilpinen, Laurence B. Lovat, Hayley J. Luxton & Hayley C. Whitaker
Division of Cancer Genomics, National Cancer Center Research Institute, National Cancer Center, Tokyo, Japan
Yasuhito Arai, Natsuko Hama, Fumie Hosoda, Hiromi Nakamura, Tatsuhiro Shibata, Yasushi Totoki & Shinichi Yachida
DLR Project Management Agency, Bonn, Germany
Axel Aretz
Tokyo Women’s Medical University, Tokyo, Japan
Shun-ichi Ariizumi & Masakazu Yamamoto
Center for Molecular Oncology, Memorial Sloan Kettering Cancer Center, New York, NY, USA
Joshua Armenia, Hsiao-Wei Chen, Jianjiong Gao, Ritika Kundra, Francisco Sanchez-Vega, Nikolaus Schultz & Hongxin Zhang
Los Alamos National Laboratory, Los Alamos, NM, USA
Laurent Arnould
Department of Pathology, University Health Network, Toronto General Hospital, Toronto, ON, Canada
Sylvia Asa, Michael H. A. Roehrl & Theodorus Van der Kwast
Nottingham University Hospitals NHS Trust, Nottingham, UK
Sylvia Asa, Simon L. Parsons & Ming Tsao
Epigenomics and Cancer Risk Factors, German Cancer Research Center (DKFZ), Heidelberg, Germany
Yassen Assenov
Computational Biology Program, Ontario Institute for Cancer Research, Toronto, ON, Canada
Gurnit Atwal, Philip Awadalla, Jonathan Barenboim, Vinayak Bhandari, Ivan Borozan, Paul C. Boutros, Lewis Jonathan Dursi, Shadrielle M. G. Espiritu, Natalie S. Fox, Michael Fraser, Syed Haider, Vincent Huang, Keren Isaev, Wei Jiao, Christopher M. Lalansingh, Emilie Lalonde, Fabien C. Lamaze, Constance H. Li, Julie Livingstone, Christine P’ng, Marta Paczkowska, Stephenie D. Prokopec, Jüri Reimand, Veronica Y. Sabelnykova, Adriana Salcedo, Yu-Jia Shiah, Solomon I. Shorser, Shimin Shuai, Jared T. Simpson, Lincoln D. Stein, Ren X. Sun, Lina Wadi, Gavin W. Wilson, Adam J. Wright, Takafumi N. Yamaguchi, Fouad Yousif & Denis Yuen
Department of Molecular Genetics, University of Toronto, Toronto, ON, Canada
Gurnit Atwal, Philip Awadalla, Gary D. Bader, Shimin Shuai & Lincoln D. Stein
Vector Institute, Toronto, ON, Canada
Gurnit Atwal, Quaid D. Morris, Yulia Rubanova & Jeffrey A. Wintersinger
Hematopathology Section, Institute of Pathology, Christian-Albrechts-University, Kiel, Germany
Sietse Aukema, Wolfram Klapper, Julia Richter & Monika Szczepanowski
Department of Pathology and Laboratory Medicine, School of Medicine, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
J. Todd Auman & Charles M. Perou
Department of Cancer Genetics, Institute for Cancer Research, Oslo University Hospital, The Norwegian Radium Hospital, Oslo, Norway
Miriam R. R. Aure, Anne-Lise Børresen-Dale & Anita Langerød
Pathology, Hospital Clinic, Institut d’Investigacions Biomèdiques August Pi i Sunyer (IDIBAPS), University of Barcelona, Barcelona, Spain
Marta Aymerich
Department of Veterinary Medicine, Transmissible Cancer Group, University of Cambridge, Cambridge, UK
Adrian Baez-Ortega
Alvin J. Siteman Cancer Center, Washington University School of Medicine, St. Louis, MO, USA
Matthew H. Bailey, Li Ding, Robert S. Fulton, Ramaswamy Govindan & Michael D. McLellan
Wolfson Wohl Cancer Research Centre, Institute of Cancer Sciences, University of Glasgow, Glasgow, UK
Peter J. Bailey, Andrew V. Biankin, David K. Chang, Susanna L. Cooke, Fraser R. Duthie, Janet S. Graham, Nigel B. Jamieson, Elizabeth A. Musgrove & Derek W. Wright
Lineberger Comprehensive Cancer Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Saianand Balu, Tom Bodenheimer, D. Neil Hayes, Austin J. Hepperla, Katherine A. Hoadley, Alan P. Hoyle, Stuart R. Jefferys, Shaowu Meng, Lisle E. Mose, Grant Sanders, Yan Shi, Janae V. Simons & Matthew G. Soloway
Broad Institute of MIT and Harvard, Cambridge, MA, USA
Pratiti Bandopadhayay, Rameen Beroukhim, Angela N. Brooks, Susan Bullman, John Busanovich, Andrew D. Cherniack, Juok Cho, Carrie Cibulskis, Kristian Cibulskis, David Craft, Timothy Defreitas, Andrew J. Dunford, Scott Frazer, Stacey B. Gabriel, Nils Gehlenborg, Gad Getz, Manaswi Gupta, Gavin Ha, Nicholas J. Haradhvala, David I. Heiman, Julian M. Hess, Manolis Kellis, Jaegil Kim, Kiran Kumar, Kirsten Kübler, Eric Lander, Michael S. Lawrence, Ignaty Leshchiner, Pei Lin, Ziao Lin, Dimitri Livitz, Yosef E. Maruvka, Samuel R. Meier, Matthew Meyerson, Michael S. Noble, Chandra Sekhar Pedamallu, Paz Polak, Esther Rheinbay, Daniel Rosebrock, Mara Rosenberg, Gordon Saksena, Richard Sallari, Steven E. Schumacher, Ayellet V. Segre, Ofer Shapira, Juliann Shih, Nasa Sinnott-Armstrong, Oliver Spiro, Chip Stewart, Amaro Taylor-Weiner, Grace Tiao, Douglas Voet, Jeremiah A. Wala, Cheng-Zhong Zhang & Hailei Zhang
Dana-Farber/Boston Children’s Cancer and Blood Disorders Center, Boston, MA, USA
Pratiti Bandopadhayay
Department of Pediatrics, Harvard Medical School, Boston, MA, USA
Pratiti Bandopadhayay
Leeds Institute of Medical Research @ St. James’s, University of Leeds, St. James’s University Hospital, Leeds, UK
Rosamonde E. Banks & Naveen Vasudev
Department of Pathology and Diagnostics, University and Hospital Trust of Verona, Verona, Italy
Stefano Barbi, Vincenzo Corbo & Michele Simbolo
Department of Surgery, Princess Alexandra Hospital, Brisbane, QLD, Australia
Andrew P. Barbour
Surgical Oncology Group, Diamantina Institute, University of Queensland, Brisbane, QLD, Australia
Andrew P. Barbour
Department of Population and Quantitative Health Sciences, Case Western Reserve University School of Medicine, Cleveland, OH, USA
Jill Barnholtz-Sloan
Research Health Analytics and Informatics, University Hospitals Cleveland Medical Center, Cleveland, OH, USA
Jill Barnholtz-Sloan
Gloucester Royal Hospital, Gloucester, UK
Hugh Barr
European Molecular Biology Laboratory, European Bioinformatics Institute (EMBL-EBI), Cambridge, UK
Elisabet Barrera, Wojciech Bazant, Ewan Birney, Rich Boyce, Alvis Brazma, Andy Cafferkey, Claudia Calabrese, Paul Flicek, Nuno A. Fonseca, Anja Füllgrabe, Moritz Gerstung, Santiago Gonzalez, Liliana Greger, Maria Keays, Jan O. Korbel, Alfonso Muñoz, Steven J. Newhouse, David Ocana, Irene Papatheodorou, Robert Petryszak, Roland F. Schwarz, Charles Short, Oliver Stegle & Lara Urban
Diagnostic Development, Ontario Institute for Cancer Research, Toronto, ON, Canada
John Bartlett & Ilinca Lungu
Barcelona Supercomputing Center (BSC), Barcelona, Spain
Javier Bartolome, Mattia Bosio, Ana Dueso-Barroso, J. Lynn Fink, Josep L. L. Gelpi, Ana Milovanovic, Montserrat Puiggròs, Javier Bartolomé Rodriguez, Romina Royo, David Torrents, Alfonso Valencia, Miguel Vazquez, David Vicente & Izar Villasante
Arnie Charbonneau Cancer Institute, University of Calgary, Calgary, AB, Canada
Oliver F. Bathe
Departments of Surgery and Oncology, University of Calgary, Calgary, AB, Canada
Oliver F. Bathe
Department of Pathology, Oslo University Hospital, The Norwegian Radium Hospital, Oslo, Norway
Daniel Baumhoer & Bodil Bjerkehagen
PanCuRx Translational Research Initiative, Ontario Institute for Cancer Research, Toronto, ON, Canada
Prashant Bavi, Michelle Chan-Seng-Yue, Sean Cleary, Robert E. Denroche, Steven Gallinger, Robert C. Grant, Gun Ho Jang, Sangeetha Kalimuthu, Ilinca Lungu, John D. McPherson, Faiyaz Notta, Michael H. A. Roehrl, Gavin W. Wilson & Julie M. Wilson
Department of Oncology, Sidney Kimmel Comprehensive Cancer Center at Johns Hopkins University School of Medicine, Baltimore, MD, USA
Stephen B. Baylin, Nilanjan Chatterjee, Leslie Cope, Ludmila Danilova & Ralph H. Hruban
University Hospital Southampton NHS Foundation Trust, Southampton, UK
Stephen B. Baylin & Tim Dudderidge
Royal Stoke University Hospital, Stoke-on-Trent, UK
Duncan Beardsmore & Christopher Umbricht
Genome Sequence Informatics, Ontario Institute for Cancer Research, Toronto, ON, Canada
Timothy A. Beck, Bob Gibson, Lawrence E. Heisler, Xuemei Luo & Morgan L. Taschuk
Human Longevity Inc, San Diego, CA, USA
Timothy A. Beck
Olivia Newton-John Cancer Research Institute, La Trobe University, Heidelberg, VIC, Australia
Andreas Behren & Jonathan Cebon
Computer Network Information Center, Chinese Academy of Sciences, Beijing, China
Beifang Niu
Genome Canada, Ottawa, ON, Canada
Cindy Bell
CNAG-CRG, Centre for Genomic Regulation (CRG), Barcelona Institute of Science and Technology (BIST), Barcelona, Spain
Sergi Beltran, Ivo G. Gut, Marta Gut, Simon C. Heath, Tomas Marques-Bonet, Arcadi Navarro, Miranda D. Stobbe, Jean-Rémi Trotta & Justin P. Whalley
Universitat Pompeu Fabra (UPF), Barcelona, Spain
Sergi Beltran, Mattia Bosio, German M. Demidov, Oliver Drechsel, Ivo G. Gut, Marta Gut, Simon C. Heath, Francesc Muyas, Stephan Ossowski, Aparna Prasad, Raquel Rabionet, Miranda D. Stobbe & Hana Susak
Buck Institute for Research on Aging, Novato, CA, USA
Christopher Benz & Christina Yau
Duke University Medical Center, Durham, NC, USA
Andrew Berchuck
Department of Human Genetics, Hannover Medical School, Hannover, Germany
Anke K. Bergmann
Center for Bioinformatics and Functional Genomics, Cedars-Sinai Medical Center, Los Angeles, CA, USA
Benjamin P. Berman & Huy Q. Dinh
Department of Biomedical Sciences, Cedars-Sinai Medical Center, Los Angeles, CA, USA
Benjamin P. Berman
The Hebrew University Faculty of Medicine, Jerusalem, Israel
Benjamin P. Berman
Barts Cancer Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London, UK
Daniel M. Berney & Yong-Jie Lu
Department of Computer Science, Bioinformatics Group, University of Leipzig, Leipzig, Germany
Stephan H. Bernhart, Hans Binder, Steve Hoffmann & Peter F. Stadler
Interdisciplinary Center for Bioinformatics, University of Leipzig, Leipzig, Germany
Stephan H. Bernhart, Hans Binder, Steve Hoffmann, Helene Kretzmer & Peter F. Stadler
Transcriptome Bioinformatics, LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
Stephan H. Bernhart, Steve Hoffmann, Helene Kretzmer & Peter F. Stadler
Department of Medical Oncology, Dana-Farber Cancer Institute, Boston, MA, USA
Rameen Beroukhim, Angela N. Brooks, Susan Bullman, Andrew D. Cherniack, Levi Garraway, Matthew Meyerson, Chandra Sekhar Pedamallu, Steven E. Schumacher, Juliann Shih & Jeremiah A. Wala
Department of Cancer Biology, Dana-Farber Cancer Institute, Boston, MA, USA
Rameen Beroukhim, Aquila Fatima, Andrea L. Richardson, Steven E. Schumacher, Ofer Shapira, Andrew Tutt & Jeremiah A. Wala
Harvard Medical School, Boston, MA, USA
Rameen Beroukhim, Gad Getz, Kirsten Kübler, Matthew Meyerson, Chandra Sekhar Pedamallu, Paz Polak, Esther Rheinbay & Jeremiah A. Wala
USC Norris Comprehensive Cancer Center, University of Southern California, Los Angeles, CA, USA
Mario Berrios, Moiz S. Bootwalla, Andrea Holbrook, Phillip H. Lai, Dennis T. Maglinte, David J. Van Den Berg & Daniel J. Weisenberger
Department of Diagnostics and Public Health, University and Hospital Trust of Verona, Verona, Italy
Samantha Bersani, Ivana Cataldo, Claudio Luchini & Maria Scardoni
Department of Mathematics, Aarhus University, Aarhus, Denmark
Johanna Bertl & Asger Hobolth
Department of Molecular Medicine (MOMA), Aarhus University Hospital, Aarhus N, Denmark
Johanna Bertl, Henrik Hornshøj, Malene Juul, Randi Istrup Juul, Tobias Madsen, Morten Muhlig Nielsen & Jakob Skou Pedersen
Instituto Carlos Slim de la Salud, Mexico City, Mexico
Miguel Betancourt
Department of Medical Biophysics, University of Toronto, Toronto, ON, Canada
Vinayak Bhandari, Paul C. Boutros, Robert G. Bristow, Keren Isaev, Constance H. Li, Jüri Reimand, Michael H. A. Roehrl & Bradly G. Wouters
Cancer Division, Garvan Institute of Medical Research, Kinghorn Cancer Centre, University of New South Wales (UNSW Sydney), Sydney, NSW, Australia
Andrew V. Biankin, David K. Chang, Lorraine A. Chantrill, Angela Chou, Anthony J. Gill, Amber L. Johns, James G. Kench, David K. Miller, Adnan M. Nagrial, Marina Pajic, Mark Pinese, Ilse Rooman, Christopher J. Scarlett, Christopher W. Toon & Jianmin Wu
South Western Sydney Clinical School, Faculty of Medicine, University of New South Wales (UNSW Sydney), Liverpool, NSW, Australia
Andrew V. Biankin
West of Scotland Pancreatic Unit, Glasgow Royal Infirmary, Glasgow, UK
Andrew V. Biankin & Nigel B. Jamieson
Center for Digital Health, Berlin Institute of Health and Charitè - Universitätsmedizin Berlin, Berlin, Germany
Matthias Bieg
Heidelberg Center for Personalized Oncology (DKFZ-HIPO), German Cancer Research Center (DKFZ), Heidelberg, Germany
Matthias Bieg, Ivo Buchhalter, Barbara Hutter & Nagarajan Paramasivam
The Preston Robert Tisch Brain Tumor Center, Duke University Medical Center, Durham, NC, USA
Darell Bigner
Massachusetts General Hospital, Boston, MA, USA
Michael Birrer, Vikram Deshpande, William C. Faquin, Nicholas J. Haradhvala, Kirsten Kübler, Michael S. Lawrence, David N. Louis, Yosef E. Maruvka, G. Petur Nielsen, Esther Rheinbay, Mara Rosenberg, Dennis C. Sgroi & Chin-Lee Wu
National Institute of Biomedical Genomics, Kalyani, West Bengal, India
Nidhan K. Biswas, Arindam Maitra & Partha P. Majumder
Institute of Clinical Medicine and Institute of Oral Biology, University of Oslo, Oslo, Norway
Bodil Bjerkehagen
University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Lori Boice, Mei Huang, Sonia Puig & Leigh B. Thorne
ARC-Net Centre for Applied Research on Cancer, University and Hospital Trust of Verona, Verona, Italy
Giada Bonizzato, Cinzia Cantù, Ivana Cataldo, Vincenzo Corbo, Sonia Grimaldi, Rita T. Lawlor, Andrea Mafficini, Borislav C. Rusev, Aldo Scarpa, Katarzyna O. Sikora, Nicola Sperandio, Alain Viari & Caterina Vicentini
The Institute of Cancer Research, London, UK
Johann S. De Bono, Niedzica Camacho, Colin S. Cooper, Sandra E. Edwards, Rosalind A. Eeles, Zsofia Kote-Jarai, Daniel A. Leongamornlert, Lucy Matthews & Sue Merson
Centre for Computational Biology, Duke-NUS Medical School, Singapore, Singapore
Arnoud Boot, Ioana Cutcutache, Mi Ni Huang, John R. McPherson, Steven G. Rozen & Yang Wu
Programme in Cancer and Stem Cell Biology, Duke-NUS Medical School, Singapore, Singapore
Arnoud Boot, Ioana Cutcutache, Mi Ni Huang, John R. McPherson, Steven G. Rozen, Patrick Tan, Bin Tean Teh & Yang Wu
Division of Oncology and Pathology, Department of Clinical Sciences Lund, Lund University, Lund, Sweden
Ake Borg, Markus Ringnér & Johan Staaf
Department of Pediatric Oncology, Hematology and Clinical Immunology, Heinrich-Heine-University, Düsseldorf, Germany
Arndt Borkhardt & Jessica I. Hoell
Laboratory for Medical Science Mathematics, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan
Keith A. Boroevich, Todd A. Johnson, Michael S. Lawrence & Tatsuhiko Tsunoda
RIKEN Center for Integrative Medical Sciences, Yokohama, Japan
Keith A. Boroevich, Akihiro Fujimoto, Masashi Fujita, Mayuko Furuta, Kazuhiro Maejima, Hidewaki Nakagawa, Kaoru Nakano & Aya Sasaki-Oku
Department of Internal Medicine/Hematology, Friedrich-Ebert-Hospital, Neumünster, Germany
Christoph Borst & Siegfried Haas
Departments of Dermatology and Pathology, Yale University, New Haven, CT, USA
Marcus Bosenberg
Centre for Genomic Regulation (CRG), The Barcelona Institute of Science and Technology, Barcelona, Spain
Mattia Bosio, German M. Demidov, Oliver Drechsel, Georgia Escaramis, Xavier Estivill, Aliaksei Z. Holik, Francesc Muyas, Stephan Ossowski, Raquel Rabionet & Hana Susak
Radcliffe Department of Medicine, University of Oxford, Oxford, UK
Jacqueline Boultwood
Canadian Center for Computational Genomics, McGill University, Montreal, QC, Canada
Guillaume Bourque
Department of Human Genetics, McGill University, Montreal, QC, Canada
Guillaume Bourque, Mark Lathrop & Yasser Riazalhosseini
Department of Human Genetics, University of California Los Angeles, Los Angeles, CA, USA
Paul C. Boutros
Department of Pharmacology, University of Toronto, Toronto, ON, Canada
Paul C. Boutros
Faculty of Medicine and Health Technology, Tampere University and Tays Cancer Center, Tampere University Hospital, Tampere, Finland
G. Steven Bova & Tapio Visakorpi
Haematology, Leeds Teaching Hospitals NHS Trust, Leeds, UK
David T. Bowen
Translational Research and Innovation, Centre Léon Bérard, Lyon, France
Sandrine Boyault
Fox Chase Cancer Center, Philadelphia, PA, USA
Jeffrey Boyd & Elaine R. Mardis
International Agency for Research on Cancer, World Health Organization, Lyon, France
Paul Brennan & Ghislaine Scelo
Earlham Institute, Norwich, UK
Daniel S. Brewer & Colin S. Cooper
Norwich Medical School, University of East Anglia, Norwich, UK
Daniel S. Brewer & Colin S. Cooper
Department of Molecular Biology, Faculty of Science, Radboud Institute for Molecular Life Sciences, Radboud University, Nijmegen, HB, The Netherlands
Arie B. Brinkman
CRUK Manchester Institute and Centre, Manchester, UK
Robert G. Bristow
Department of Radiation Oncology, University of Toronto, Toronto, ON, Canada
Robert G. Bristow
Division of Cancer Sciences, Manchester Cancer Research Centre, University of Manchester, Manchester, UK
Robert G. Bristow
Radiation Medicine Program, Princess Margaret Cancer Centre, Toronto, ON, Canada
Robert G. Bristow & Fei-Fei Fei Liu
Department of Pathology, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA
Jane E. Brock & Sabina Signoretti
Department of Surgery, Division of Thoracic Surgery, The Johns Hopkins University School of Medicine, Baltimore, MD, USA
Malcolm Brock
Division of Molecular Pathology, The Netherlands Cancer Institute, Oncode Institute, Amsterdam, CX, The Netherlands
Annegien Broeks & Jos Jonkers
Department of Biomolecular Engineering, University of California Santa Cruz, Santa Cruz, CA, USA
Angela N. Brooks, David Haan, Maximillian G. Marin, Thomas J. Matthew, Yulia Newton, Cameron M. Soulette & Joshua M. Stuart
UC Santa Cruz Genomics Institute, University of California Santa Cruz, Santa Cruz, CA, USA
Angela N. Brooks, Brian Craft, Mary J. Goldman, David Haussler, Joshua M. Stuart & Jingchun Zhu
Division of Applied Bioinformatics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Benedikt Brors, Lars Feuerbach, Chen Hong, Charles David Imbusch & Lina Sieverling
German Cancer Consortium (DKTK), German Cancer Research Center (DKFZ), Heidelberg, Germany
Benedikt Brors, Barbara Hutter, Peter Lichter, Dirk Schadendorf & Holger Sültmann
National Center for Tumor Diseases (NCT) Heidelberg, Heidelberg, Germany
Benedikt Brors, Barbara Hutter, Holger Sültmann & Thorsten Zenz
Center for Biological Sequence Analysis, Department of Bio and Health Informatics, Technical University of Denmark, Lyngby, Denmark
Søren Brunak
Novo Nordisk Foundation Center for Protein Research, University of Copenhagen, Copenhagen, Denmark
Søren Brunak
Institute for Molecular Bioscience, University of Queensland, St. Lucia, Brisbane, QLD, Australia
Timothy J. C. Bruxner, Oliver Holmes, Stephen H. Kazakoff, Conrad R. Leonard, Felicity Newell, Katia Nones, Ann-Marie Patch, John V. Pearson, Michael C. Quinn, Nick M. Waddell, Nicola Waddell, Scott Wood & Qinying Xu
Biomedical Engineering, Oregon Health and Science University, Portland, OR, USA
Alex Buchanan & Kyle Ellrott
Division of Theoretical Bioinformatics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Ivo Buchhalter, Calvin Wing Yiu Chan, Roland Eils, Michael C. Heinold, Carl Herrmann, Natalie Jäger, Rolf Kabbe, Jules N. A. Kerssemakers, Kortine Kleinheinz, Nagarajan Paramasivam, Manuel Prinz, Matthias Schlesner & Johannes Werner
Institute of Pharmacy and Molecular Biotechnology and BioQuant, Heidelberg University, Heidelberg, Germany
Ivo Buchhalter, Roland Eils, Michael C. Heinold, Carl Herrmann, Daniel Hübschmann, Kortine Kleinheinz & Umut H. Toprak
Federal Ministry of Education and Research, Berlin, Germany
Christiane Buchholz
Melanoma Institute Australia, University of Sydney, Sydney, NSW, Australia
Hazel Burke, Ricardo De Paoli-Iseppi, Nicholas K. Hayward, Peter Hersey, Valerie Jakrot, Hojabr Kakavand, Georgina V. Long, Graham J. Mann, Robyn P. M. Saw, Richard A. Scolyer, Ping Shang, Andrew J. Spillane, Jonathan R. Stretch, John F. F. Thompson & James S. Wilmott
Pediatric Hematology and Oncology, University Hospital Muenster, Muenster, Germany
Birgit Burkhardt
Department of Pathology, Johns Hopkins University School of Medicine, Baltimore, MD, USA
Kathleen H. Burns & Christopher Umbricht
McKusick-Nathans Institute of Genetic Medicine, Sidney Kimmel Comprehensive Cancer Center at Johns Hopkins University School of Medicine, Baltimore, MD, USA
Kathleen H. Burns
Foundation Medicine, Inc, Cambridge, MA, USA
John Busanovich
Department of Biomedical Data Science, Stanford University School of Medicine, Stanford, CA, USA
Carlos D. Bustamante & Francisco M. De La Vega
Department of Genetics, Stanford University School of Medicine, Stanford, CA, USA
Carlos D. Bustamante, Francisco M. De La Vega, Suyash S. Shringarpure, Nasa Sinnott-Armstrong & Mark H. Wright
Bakar Computational Health Sciences Institute and Department of Pediatrics, University of California, San Francisco, CA, USA
Atul J. Butte & Jieming Chen
Institute of Clinical Medicine, Faculty of Medicine, University of Oslo, Oslo, Norway
Anne-Lise Børresen-Dale
National Cancer Institute, National Institutes of Health, Bethesda, MD, USA
Samantha J. Caesar-Johnson, John A. Demchok, Ina Felau, Roy Tarnuzzer, Zhining Wang, Liming Yang, Jean C. Zenklusen & Jiashan Zhang
Royal Marsden NHS Foundation Trust, London and Sutton, UK
Declan Cahill, Nening M. Dennis, Tim Dudderidge, Rosalind A. Eeles, Cyril Fisher, Steven Hazell, Vincent Khoo, Pardeep Kumar, Naomi Livni, Erik Mayer, David Nicol, Christopher Ogden, Edward W. Rowe, Sarah Thomas, Alan Thompson & Nicholas van As
Genome Biology Unit, European Molecular Biology Laboratory (EMBL), Heidelberg, Germany
Claudia Calabrese, Serap Erkek, Moritz Gerstung, Santiago Gonzalez, Nina Habermann, Wolfgang Huber, Lara Jerman, Jan O. Korbel, Esa Pitkänen, Benjamin Raeder, Tobias Rausch, Vasilisa A. Rudneva, Oliver Stegle, Stephanie Sungalee, Lara Urban, Sebastian M. Waszak, Joachim Weischenfeldt & Sergei Yakneen
Department of Oncology, University of Cambridge, Cambridge, UK
Carlos Caldas & Suet-Feung Chin
Li Ka Shing Centre, Cancer Research UK Cambridge Institute, University of Cambridge, Cambridge, UK
Carlos Caldas, Suet-Feung Chin, Ruben M. Drews, Paul A. Edwards, Matthew Eldridge, Steve Hawkins, Andy G. Lynch, Geoff Macintyre, Florian Markowetz, Charlie E. Massie, David E. Neal, Simon Tavaré & Ke Yuan
Institut Gustave Roussy, Villejuif, France
Fabien Calvo
Cambridge University Hospitals NHS Foundation Trust, Cambridge, UK
Peter J. Campbell, Vincent J. Gnanapragasam, William Howat, Thomas J. Mitchell, David E. Neal, Nimish C. Shah & Anne Y. Warren
Department of Haematology, University of Cambridge, Cambridge, UK
Peter J. Campbell
Anatomia Patológica, Hospital Clinic, Institut d’Investigacions Biomèdiques August Pi i Sunyer (IDIBAPS), University of Barcelona, Barcelona, Spain
Elias Campo
Spanish Ministry of Science and Innovation, Madrid, Spain
Elias Campo
University of Michigan Comprehensive Cancer Center, Ann Arbor, MI, USA
Thomas E. Carey
Department for BioMedical Research, University of Bern, Bern, Switzerland
Joana Carlevaro-Fita
Department of Medical Oncology, Inselspital, University Hospital and University of Bern, Bern, Switzerland
Joana Carlevaro-Fita, Rory Johnson & Andrés Lanzós
Graduate School for Cellular and Biomedical Sciences, University of Bern, Bern, Switzerland
Joana Carlevaro-Fita & Andrés Lanzós
University of Pavia, Pavia, Italy
Mario Cazzola & Luca Malcovati
University of Alabama at Birmingham, Birmingham, AL, USA
Robert Cerfolio
UHN Program in BioSpecimen Sciences, Toronto General Hospital, Toronto, ON, Canada
Dianne E. Chadwick, Sheng-Ben Liang, Michael H. A. Roehrl & Sagedeh Shahabi
Department of Urology, Icahn School of Medicine at Mount Sinai, New York, NY, USA
Dimple Chakravarty
Centre for Law and Genetics, University of Tasmania, Sandy Bay Campus, Hobart, TAS, Australia
Don Chalmers
Faculty of Biosciences, Heidelberg University, Heidelberg, Germany
Calvin Wing Yiu Chan, Chen Hong & Lina Sieverling
Department of Biochemistry, Microbiology and Immunology, Faculty of Medicine, University of Ottawa, Ottawa, ON, Canada
Kin Chan
Division of Anatomic Pathology, Mayo Clinic, Rochester, MN, USA
Vishal S. Chandan
Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, MD, USA
Stephen J. Chanock, Xing Hua, Lisa Mirabello, Lei Song & Bin Zhu
Illawarra Shoalhaven Local Health District L3 Illawarra Cancer Care Centre, Wollongong Hospital, Wollongong, NSW, Australia
Lorraine A. Chantrill
BioForA, French National Institute for Agriculture, Food, and Environment (INRAE), ONF, Orléans, France
Aurélien Chateigner
Department of Biostatistics, Bloomberg School of Public Health, Johns Hopkins University, Baltimore, MD, USA
Nilanjan Chatterjee
University of California San Diego, San Diego, CA, USA
Zhaohong Chen, Michelle T. Dow, Claudiu Farcas, S. M. Ashiqul Islam, Antonios Koures, Lucila Ohno-Machado, Christos Sotiriou & Ashley Williams
Division of Experimental Pathology, Mayo Clinic, Rochester, MN, USA
Jeremy Chien
Centre for Cancer Research, The Westmead Institute for Medical Research, University of Sydney, Sydney, NSW, Australia
Yoke-Eng Chiew, Angela Chou, Jillian A. Hung, Catherine J. Kennedy, Graham J. Mann, Gulietta M. Pupo, Sarah-Jane Schramm, Varsha Tembe & Anna deFazio
Department of Gynaecological Oncology, Westmead Hospital, Sydney, NSW, Australia
Yoke-Eng Chiew, Jillian A. Hung, Catherine J. Kennedy & Anna deFazio
PDXen Biosystems Inc, Seoul, South Korea
Sunghoon Cho
Korea Advanced Institute of Science and Technology, Daejeon, South Korea
Jung Kyoon Choi, Young Seok Ju & Christopher J. Yoon
Electronics and Telecommunications Research Institute, Daejeon, South Korea
Wan Choi, Seung-Hyup Jeon, Hyunghwan Kim & Youngchoon Woo
Institut National du Cancer (INCA), Boulogne-Billancourt, France
Christine Chomienne & Iris Pauporté
Department of Genetics, Informatics Institute, University of Alabama at Birmingham, Birmingham, AL, USA
Zechen Chong
Division of Medical Oncology, National Cancer Centre, Singapore, Singapore
Su Pin Choo
Medical Oncology, University and Hospital Trust of Verona, Verona, Italy
Sara Cingarlini & Michele Milella
Department of Pediatrics, University Hospital Schleswig-Holstein, Kiel, Germany
Alexander Claviez
Hepatobiliary/Pancreatic Surgical Oncology Program, University Health Network, Toronto, ON, Canada
Sean Cleary, Ashton A. Connor & Steven Gallinger
School of Biological Sciences, University of Auckland, Auckland, New Zealand
Nicole Cloonan
Department of Surgery, University of Melbourne, Parkville, VIC, Australia
Marek Cmero
The Murdoch Children’s Research Institute, Royal Children’s Hospital, Parkville, VIC, Australia
Marek Cmero
Walter and Eliza Hall Institute, Parkville, VIC, Australia
Marek Cmero
Vancouver Prostate Centre, Vancouver, Canada
Colin C. Collins, Nilgun Donmez, Faraz Hach, Salem Malikic, S. Cenk Sahinalp, Iman Sarrafi & Raunak Shrestha
Lunenfeld-Tanenbaum Research Institute, Mount Sinai Hospital, Toronto, ON, Canada
Ashton A. Connor, Steven Gallinger, Robert C. Grant, Treasa A. McPherson & Iris Selander
University of East Anglia, Norwich, UK
Colin S. Cooper
Norfolk and Norwich University Hospital NHS Trust, Norwich, UK
Matthew G. Cordes, Catrina C. Fronick & Tom Roques
Victorian Institute of Forensic Medicine, Southbank, VIC, Australia
Stephen M. Cordner
Department of Biomedical Informatics, Harvard Medical School, Boston, MA, USA
Isidro Cortés-Ciriano, Jake June-Koo Lee & Peter J. Park
Department of Chemistry, Centre for Molecular Science Informatics, University of Cambridge, Cambridge, UK
Isidro Cortés-Ciriano
Ludwig Center at Harvard Medical School, Boston, MA, USA
Isidro Cortés-Ciriano, Jake June-Koo Lee & Peter J. Park
Human Genome Sequencing Center, Baylor College of Medicine, Houston, TX, USA
Kyle Covington, HarshaVardhan Doddapaneni, Richard A. Gibbs, Jianhong Hu, Joy C. Jayaseelan, Viktoriya Korchina, Lora Lewis, Donna M. Muzny, Linghua Wang, David A. Wheeler & Liu Xi
Peter MacCallum Cancer Centre, University of Melbourne, Melbourne, VIC, Australia
Prue A. Cowin, Anne Hamilton, Gisela Mir Arnau & Ravikiran Vedururu
Physics Division, Optimization and Systems Biology Lab, Massachusetts General Hospital, Boston, MA, USA
David Craft
Department of Medicine, Baylor College of Medicine, Houston, TX, USA
Chad J. Creighton
University of Cologne, Cologne, Germany
Yupeng Cun, Martin Peifer & Tsun-Po Yang
International Genomics Consortium, Phoenix, AZ, USA
Erin Curley & Troy Shelton
Genomics Research Program, Ontario Institute for Cancer Research, Toronto, ON, Canada
Karolina Czajka, Jenna Eagles, Thomas J. Hudson, Jeremy Johns, Faridah Mbabaali, John D. McPherson, Jessica K. Miller, Danielle Pasternack, Michelle Sam & Lee E. Timms
Barking Havering and Redbridge University Hospitals NHS Trust, Romford, UK
Bogdan Czerniak, Adel El-Naggar & David Khoo
Children’s Hospital at Westmead, University of Sydney, Sydney, NSW, Australia
Rebecca A. Dagg
Department of Medicine, Section of Endocrinology, University and Hospital Trust of Verona, Verona, Italy
Maria Vittoria Davi
Computational Biology Center, Memorial Sloan Kettering Cancer Center, New York, NY, USA
Natalie R. Davidson, Andre Kahles, Kjong-Van Lehmann, Alessandro Pastore, Gunnar Rätsch, Chris Sander, Yasin Senbabaoglu & Nicholas D. Socci
Department of Biology, ETH Zurich, Zürich, Switzerland
Natalie R. Davidson, Andre Kahles, Kjong-Van Lehmann, Gunnar Rätsch & Stefan G. Stark
Department of Computer Science, ETH Zurich, Zurich, Switzerland
Natalie R. Davidson, Andre Kahles, Kjong-Van Lehmann & Gunnar Rätsch
SIB Swiss Institute of Bioinformatics, Lausanne, Switzerland
Natalie R. Davidson, Andre Kahles, Kjong-Van Lehmann, Gunnar Rätsch & Stefan G. Stark
Weill Cornell Medical College, New York, NY, USA
Natalie R. Davidson, Bishoy M. Faltas & Gunnar Rätsch
Academic Department of Medical Genetics, University of Cambridge, Addenbrooke’s Hospital, Cambridge, UK
Helen Davies & Serena Nik-Zainal
MRC Cancer Unit, University of Cambridge, Cambridge, UK
Helen Davies, Rebecca C. Fitzgerald, Nicola Grehan, Serena Nik-Zainal & Maria O’Donovan
Departments of Pediatrics and Genetics, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Ian J. Davis
Seven Bridges Genomics, Charlestown, MA, USA
Brandi N. Davis-Dusenbery, Sinisa Ivkovic, Milena Kovacevic, Ana Mijalkovic Lazic, Sanja Mijalkovic, Mia Nastic, Petar Radovic & Nebojsa Tijanic
Annai Systems, Inc, Carlsbad, CA, USA
Francisco M. De La Vega, Tal Shmaya & Dai-Ying Wu
Department of Pathology, General Hospital of Treviso, Department of Medicine, University of Padua, Treviso, Italy
Angelo P. Dei Tos
Department of Computational Biology, University of Lausanne, Lausanne, Switzerland
Olivier Delaneau
Department of Genetic Medicine and Development, University of Geneva Medical School, Geneva, CH, Switzerland
Olivier Delaneau
Swiss Institute of Bioinformatics, University of Geneva, Geneva, CH, Switzerland
Olivier Delaneau
The Francis Crick Institute, London, UK
Jonas Demeulemeester, Stefan C. Dentro, Matthew W. Fittall, Kerstin Haase, Clemency Jolly, Maxime Tarabichi & Peter Van Loo
University of Leuven, Leuven, Belgium
Jonas Demeulemeester & Peter Van Loo
Institute of Medical Genetics and Applied Genomics, University of Tübingen, Tübingen, Germany
German M. Demidov, Francesc Muyas & Stephan Ossowski
Computational and Systems Biology, Genome Institute of Singapore, Singapore, Singapore
Deniz Demircioğlu & Jonathan Göke
School of Computing, National University of Singapore, Singapore, Singapore
Deniz Demircioğlu
Big Data Institute, Li Ka Shing Centre, University of Oxford, Oxford, UK
Stefan C. Dentro & David C. Wedge
Biomedical Data Science Laboratory, Francis Crick Institute, London, UK
Nikita Desai
Bioinformatics Group, Department of Computer Science, University College London, London, UK
Nikita Desai
The Edward S. Rogers Sr. Department of Electrical and Computer Engineering, University of Toronto, Toronto, ON, Canada
Amit G. Deshwar
Breast Cancer Translational Research Laboratory JC Heuson, Institut Jules Bordet, Brussels, Belgium
Christine Desmedt
Department of Oncology, Laboratory for Translational Breast Cancer Research, KU Leuven, Leuven, Belgium
Christine Desmedt
Institute for Research in Biomedicine (IRB Barcelona), The Barcelona Institute of Science and Technology, Barcelona, Spain
Jordi Deu-Pons, Joan Frigola, Abel Gonzalez-Perez, Ferran Muiños, Loris Mularoni, Oriol Pich, Iker Reyes-Salazar, Carlota Rubio-Perez, Radhakrishnan Sabarinathan & David Tamborero
Research Program on Biomedical Informatics, Universitat Pompeu Fabra, Barcelona, Spain
Jordi Deu-Pons, Abel Gonzalez-Perez, Ferran Muiños, Loris Mularoni, Oriol Pich, Carlota Rubio-Perez, Radhakrishnan Sabarinathan & David Tamborero
Division of Medical Oncology, Princess Margaret Cancer Centre, Toronto, ON, Canada
Neesha C. Dhani, David Hedley & Malcolm J. Moore
Department of Physiology and Biophysics, Weill Cornell Medicine, New York, NY, USA
Priyanka Dhingra, Ekta Khurana, Eric Minwei Liu & Alexander Martinez-Fundichely
Institute for Computational Biomedicine, Weill Cornell Medicine, New York, NY, USA
Priyanka Dhingra, Ekta Khurana, Eric Minwei Liu & Alexander Martinez-Fundichely
Department of Pathology, UPMC Shadyside, Pittsburgh, PA, USA
Rajiv Dhir
Independent Consultant, Wellesley, USA
Anthony DiBiase
Department of Cell and Molecular Biology, Science for Life Laboratory, Uppsala University, Uppsala, Sweden
Klev Diamanti, Jan Komorowski & Husen M. Umer
Department of Medicine and Department of Genetics, Washington University School of Medicine, St. Louis, St. Louis, MO, USA
Li Ding, Robert S. Fulton, Michael D. McLellan, Michael C. Wendl & Venkata D. Yellapantula
Hefei University of Technology, Anhui, China
Shuai Ding & Shanlin Yang
Translational Cancer Research Unit, GZA Hospitals St.-Augustinus, Center for Oncological Research, Faculty of Medicine and Health Sciences, University of Antwerp, Antwerp, Belgium
Luc Dirix, Steven Van Laere, Gert G. Van den Eynden & Peter Vermeulen
Simon Fraser University, Burnaby, BC, Canada
Nilgun Donmez, Ermin Hodzic, Salem Malikic, S. Cenk Sahinalp & Iman Sarrafi
University of Pennsylvania, Philadelphia, PA, USA
Ronny Drapkin
Faculty of Science and Technology, University of Vic—Central University of Catalonia (UVic-UCC), Vic, Spain
Ana Dueso-Barroso
The Wellcome Trust, London, UK
Michael Dunn
The Hospital for Sick Children, Toronto, ON, Canada
Lewis Jonathan Dursi
Department of Pathology, Queen Elizabeth University Hospital, Glasgow, UK
Fraser R. Duthie
Department of Genetics and Computational Biology, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia
Ken Dutton-Regester, Nicholas K. Hayward, Oliver Holmes, Peter A. Johansson, Stephen H. Kazakoff, Conrad R. Leonard, Felicity Newell, Katia Nones, Ann-Marie Patch, John V. Pearson, Antonia L. Pritchard, Michael C. Quinn, Paresh Vyas, Nicola Waddell, Scott Wood & Qinying Xu
Department of Oncology, Centre for Cancer Genetic Epidemiology, University of Cambridge, Cambridge, UK
Douglas F. Easton
Department of Public Health and Primary Care, Centre for Cancer Genetic Epidemiology, University of Cambridge, Cambridge, UK
Douglas F. Easton
Prostate Cancer Canada, Toronto, ON, Canada
Stuart Edmonds
University of Cambridge, Cambridge, UK
Paul A. Edwards, Anthony R. Green, Andy G. Lynch, Florian Markowetz & Thomas J. Mitchell
Department of Laboratory Medicine, Translational Cancer Research, Lund University Cancer Center at Medicon Village, Lund University, Lund, Sweden
Anna Ehinger
Heidelberg University, Heidelberg, Germany
Juergen Eils, Roland Eils & Daniel Hübschmann
New BIH Digital Health Center, Berlin Institute of Health (BIH) and Charité - Universitätsmedizin Berlin, Berlin, Germany
Juergen Eils, Roland Eils & Chris Lawerenz
CIBER Epidemiología y Salud Pública (CIBERESP), Madrid, Spain
Georgia Escaramis
Research Group on Statistics, Econometrics and Health (GRECS), UdG, Barcelona, Spain
Georgia Escaramis
Quantitative Genomics Laboratories (qGenomics), Barcelona, Spain
Xavier Estivill
Icelandic Cancer Registry, Icelandic Cancer Society, Reykjavik, Iceland
Jorunn E. Eyfjord, Holmfridur Hilmarsdottir & Jon G. Jonasson
State Key Laboratory of Cancer Biology, and Xijing Hospital of Digestive Diseases, Fourth Military Medical University, Shaanxi, China
Daiming Fan & Yongzhan Nie
Department of Medicine (DIMED), Surgical Pathology Unit, University of Padua, Padua, Italy
Matteo Fassan
Rigshospitalet, Copenhagen, Denmark
Francesco Favero
Center for Cancer Genomics, National Cancer Institute, National Institutes of Health, Bethesda, MD, USA
Martin L. Ferguson
Department of Biochemistry and Molecular Medicine, University of Montreal, Montreal, QC, Canada
Vincent Ferretti
Australian Institute of Tropical Health and Medicine, James Cook University, Douglas, QLD, Australia
Matthew A. Field
Department of Neuro-Oncology, Istituto Neurologico Besta, Milano, Italy
Gaetano Finocchiaro
Bioplatforms Australia, North Ryde, NSW, Australia
Anna Fitzgerald & Catherine A. Shang
Department of Pathology (Research), University College London Cancer Institute, London, UK
Adrienne M. Flanagan
Department of Surgical Oncology, Princess Margaret Cancer Centre, Toronto, ON, Canada
Neil E. Fleshner
Department of Medical Oncology, Josephine Nefkens Institute and Cancer Genomics Centre, Erasmus Medical Center, Rotterdam, CN, The Netherlands
John A. Foekens, John W. M. Martens, F. Germán Rodríguez-González, Anieta M. Sieuwerts & Marcel Smid
The University of Queensland Thoracic Research Centre, The Prince Charles Hospital, Brisbane, QLD, Australia
Kwun M. Fong
CIBIO/InBIO - Research Center in Biodiversity and Genetic Resources, Universidade do Porto, Vairão, Portugal
Nuno A. Fonseca
HCA Laboratories, London, UK
Christopher S. Foster
University of Liverpool, Liverpool, UK
Christopher S. Foster
The Azrieli Faculty of Medicine, Bar-Ilan University, Safed, Israel
Milana Frenkel-Morgenstern
Department of Neurosurgery, University of Florida, Gainesville, FL, USA
William Friedman
Department of Pathology, Graduate School of Medicine, University of Tokyo, Tokyo, Japan
Masashi Fukayama & Tetsuo Ushiku
University of Milano Bicocca, Monza, Italy
Carlo Gambacorti-Passerini
BGI-Shenzhen, Shenzhen, China
Shengjie Gao, Yong Hou, Chang Li, Lin Li, Siliang Li, Xiaobo Li, Xinyue Li, Dongbing Liu, Xingmin Liu, Qiang Pan-Hammarström, Hong Su, Jian Wang, Kui Wu, Heng Xiong, Huanming Yang, Chen Ye, Xiuqing Zhang, Yong Zhou & Shida Zhu
Department of Pathology, Oslo University Hospital Ulleval, Oslo, Norway
Øystein Garred
Center for Biomedical Informatics, Harvard Medical School, Boston, MA, USA
Nils Gehlenborg
Department Biochemistry and Molecular Biomedicine, University of Barcelona, Barcelona, Spain
Josep L. L. Gelpi
Office of Cancer Genomics, National Cancer Institute, National Institutes of Health, Bethesda, MD, USA
Daniela S. Gerhard
Cancer Epigenomics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Clarissa Gerhauser, Christoph Plass & Dieter Weichenhan
Department of Cancer Biology, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Jeffrey E. Gershenwald
Department of Surgical Oncology, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Jeffrey E. Gershenwald
Department of Computer Science, Yale University, New Haven, CT, USA
Mark Gerstein & Fabio C. P. Navarro
Department of Molecular Biophysics and Biochemistry, Yale University, New Haven, CT, USA
Mark Gerstein, Sushant Kumar, Lucas Lochovsky, Shaoke Lou, Patrick D. McGillivray, Fabio C. P. Navarro, Leonidas Salichos & Jonathan Warrell
Program in Computational Biology and Bioinformatics, Yale University, New Haven, CT, USA
Mark Gerstein, Arif O. Harmanci, Sushant Kumar, Donghoon Lee, Shantao Li, Xiaotong Li, Lucas Lochovsky, Shaoke Lou, William Meyerson, Leonidas Salichos, Jonathan Warrell, Jing Zhang & Yan Zhang
Center for Cancer Research, Massachusetts General Hospital, Boston, MA, USA
Gad Getz & Paz Polak
Department of Pathology, Massachusetts General Hospital, Boston, MA, USA
Gad Getz
Department of Pathology, Memorial Sloan Kettering Cancer Center, New York, NY, USA
Ronald Ghossein, Dilip D. Giri, Christine A. Iacobuzio-Donahue, Jorge Reis-Filho & Victor Reuter
Division of Gastroenterology and Hepatology, Mayo Clinic, Rochester, MN, USA
Nasra H. Giama, Catherine D. Moser & Lewis R. Roberts
University of Sydney, Sydney, NSW, Australia
Anthony J. Gill & James G. Kench
University of Oxford, Oxford, UK
Pelvender Gill, Freddie C. Hamdy, Katalin Karaszi, Adam Lambert, Luke Marsden, Clare Verrill & Paresh Vyas
Department of Surgery, Academic Urology Group, University of Cambridge, Cambridge, UK
Vincent J. Gnanapragasam
Department of Medicine II, University of Würzburg, Wuerzburg, Germany
Maria Elisabeth Goebler
Sylvester Comprehensive Cancer Center, University of Miami, Miami, FL, USA
Carmen Gomez
Institut Hospital del Mar d’Investigacions Mèdiques (IMIM), Barcelona, Spain
Abel Gonzalez-Perez
Genome Integrity and Structural Biology Laboratory, National Institute of Environmental Health Sciences (NIEHS), Durham, NC, USA
Dmitry A. Gordenin & Natalie Saini
St. Thomas’s Hospital, London, UK
James Gossage
Osaka International Cancer Center, Osaka, Japan
Kunihito Gotoh
Department of Pathology, Skåne University Hospital, Lund University, Lund, Sweden
Dorthe Grabau
Department of Medical Oncology, Beatson West of Scotland Cancer Centre, Glasgow, UK
Janet S. Graham
National Human Genome Research Institute, National Institutes of Health, Bethesda, MD, USA
Eric Green, Carolyn M. Hutter & Heidi J. Sofia
Centre for Cancer Research, Victorian Comprehensive Cancer Centre, University of Melbourne, Melbourne, VIC, Australia
Sean M. Grimmond
Department of Medicine, Section of Hematology/Oncology, University of Chicago, Chicago, IL, USA
Robert L. Grossman
German Center for Infection Research (DZIF), Partner Site Hamburg-Borstel-Lübeck-Riems, Hamburg, Germany
Adam Grundhoff
Bioinformatics Research Centre (BiRC), Aarhus University, Aarhus, Denmark
Qianyun Guo, Asger Hobolth & Jakob Skou Pedersen
Department of Biotechnology, Ministry of Science and Technology, Government of India, New Delhi, Delhi, India
Shailja Gupta & K. VijayRaghavan
National Cancer Centre Singapore, Singapore, Singapore
Jonathan Göke
Brandeis University, Waltham, MA, USA
James E. Haber
Department of Urologic Sciences, University of British Columbia, Vancouver, BC, Canada
Faraz Hach
Department of Internal Medicine, Stanford University, Stanford, CA, USA
Mark P. Hamilton
The University of Texas Health Science Center at Houston, Houston, TX, USA
Leng Han, Yang Yang & Xuanping Zhang
Imperial College NHS Trust, Imperial College, London, INY, UK
George B. Hanna
Senckenberg Institute of Pathology, University of Frankfurt Medical School, Frankfurt, Germany
Martin Hansmann
Department of Medicine, Division of Biomedical Informatics, UC San Diego School of Medicine, San Diego, CA, USA
Olivier Harismendy
Center for Precision Health, School of Biomedical Informatics, The University of Texas Health Science Center, Houston, TX, USA
Arif O. Harmanci
Oxford Nanopore Technologies, New York, NY, USA
Eoghan Harrington & Sissel Juul
Institute of Medical Science, University of Tokyo, Tokyo, Japan
Takanori Hasegawa, Shuto Hayashi, Seiya Imoto, Mitsuhiro Komura, Satoru Miyano, Naoki Miyoshi, Kazuhiro Ohi, Eigo Shimizu, Yuichi Shiraishi, Hiroko Tanaka & Rui Yamaguchi
Howard Hughes Medical Institute, University of California Santa Cruz, Santa Cruz, CA, USA
David Haussler
Wakayama Medical University, Wakayama, Japan
Shinya Hayami, Masaki Ueno & Hiroki Yamaue
Department of Internal Medicine, Division of Medical Oncology, Lineberger Comprehensive Cancer Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
D. Neil Hayes
University of Tennessee Health Science Center for Cancer Research, Memphis, TN, USA
D. Neil Hayes
Department of Histopathology, Salford Royal NHS Foundation Trust, Salford, UK
Stephen J. Hayes
Faculty of Biology, Medicine and Health, University of Manchester, Manchester, UK
Stephen J. Hayes
BIOPIC, ICG and College of Life Sciences, Peking University, Beijing, China
Yao He & Zemin Zhang
Peking-Tsinghua Center for Life Sciences, Peking University, Beijing, China
Yao He & Zemin Zhang
Children’s Hospital of Philadelphia, Philadelphia, PA, USA
Allison P. Heath
Department of Bioinformatics and Computational Biology and Department of Systems Biology, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Apurva M. Hegde, Yiling Lu & John N. Weinstein
Karolinska Institute, Stockholm, Sweden
Eva Hellstrom-Lindberg & Jesper Lagergren
The Donnelly Centre, University of Toronto, Toronto, ON, Canada
Mohamed Helmy & Jeffrey A. Wintersinger
Department of Medical Genetics, College of Medicine, Hallym University, Chuncheon, South Korea
Seong Gu Heo, Eun Pyo Hong & Ji Wan Park
Department of Experimental and Health Sciences, Institute of Evolutionary Biology (UPF-CSIC), Universitat Pompeu Fabra, Barcelona, Spain
José María Heredia-Genestar, Tomas Marques-Bonet & Arcadi Navarro
Health Data Science Unit, University Clinics, Heidelberg, Germany
Carl Herrmann
Massachusetts General Hospital Center for Cancer Research, Charlestown, MA, USA
Julian M. Hess & Yosef E. Maruvka
Hokkaido University, Sapporo, Japan
Satoshi Hirano & Toru Nakamura
Department of Pathology and Clinical Laboratory, National Cancer Center Hospital, Tokyo, Japan
Nobuyoshi Hiraoka
Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Katherine A. Hoadley & Tara J. Skelly
Computational Biology, Leibniz Institute on Aging - Fritz Lipmann Institute (FLI), Jena, Germany
Steve Hoffmann
University of Melbourne Centre for Cancer Research, Melbourne, VIC, Australia
Oliver Hofmann
University of Nebraska Medical Center, Omaha, NE, USA
Michael A. Hollingsworth & Sarah P. Thayer
Syntekabio Inc, Daejeon, South Korea
Jongwhi H. Hong
Department of Pathology, Academic Medical Center, Amsterdam, AZ, The Netherlands
Gerrit K. Hooijer
China National GeneBank-Shenzhen, Shenzhen, China
Yong Hou, Chang Li, Siliang Li, Xiaobo Li, Dongbing Liu, Xingmin Liu, Henk G. Stunnenberg, Hong Su, Kui Wu, Heng Xiong, Chen Ye & Shida Zhu
Division of Molecular Genetics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Volker Hovestadt, Murat Iskar, Peter Lichter, Bernhard Radlwimmer & Marc Zapatka
Division of Life Science and Applied Genomics Center, Hong Kong University of Science and Technology, Clear Water Bay, Hong Kong, China
Taobo Hu, Yogesh Kumar, Eric Z. Ma, Zhenggang Wu & Hong Xue
Icahn School of Medicine at Mount Sinai, New York, NY, USA
Kuan-lin Huang
Geneplus-Shenzhen, Shenzhen, China
Yi Huang
School of Computer Science and Technology, Xi’an Jiaotong University, Xi’an, China
Yi Huang, Jiayin Wang, Xiao Xiao & Xuanping Zhang
AbbVie, North Chicago, IL, USA
Thomas J. Hudson
Institute of Pathology, Charité – University Medicine Berlin, Berlin, Germany
Michael Hummel & Dido Lenze
Centre for Translational and Applied Genomics, British Columbia Cancer Agency, Vancouver, BC, Canada
David Huntsman
Edinburgh Royal Infirmary, Edinburgh, UK
Ted R. Hupp
Berlin Institute for Medical Systems Biology, Max Delbrück Center for Molecular Medicine, Berlin, Germany
Matthew R. Huska, Julia Markowski & Roland F. Schwarz
Department of Pediatric Immunology, Hematology and Oncology, University Hospital, Heidelberg, Germany
Daniel Hübschmann
German Cancer Research Center (DKFZ), Heidelberg, Germany
Daniel Hübschmann, Christof von Kalle & Roland F. Schwarz
Heidelberg Institute for Stem Cell Technology and Experimental Medicine (HI-STEM), Heidelberg, Germany
Daniel Hübschmann
Institute for Computational Biomedicine, Weill Cornell Medical College, New York, NY, USA
Marcin Imielinski
New York Genome Center, New York, NY, USA
Marcin Imielinski & Xiaotong Yao
Department of Urology, James Buchanan Brady Urological Institute, Johns Hopkins University School of Medicine, Baltimore, MD, USA
William B. Isaacs
Department of Preventive Medicine, Graduate School of Medicine, The University of Tokyo, Tokyo, Japan
Shumpei Ishikawa, Hiroto Katoh & Daisuke Komura
Department of Molecular and Cellular Biology, Baylor College of Medicine, Houston, TX, USA
Michael Ittmann
Department of Pathology and Immunology, Baylor College of Medicine, Houston, TX, USA
Michael Ittmann
Michael E. DeBakey Veterans Affairs Medical Center, Houston, TX, USA
Michael Ittmann
Technical University of Denmark, Lyngby, Denmark
Jose M. G. Izarzugaza
Department of Pathology, College of Medicine, Hanyang University, Seoul, South Korea
Jocelyne Jacquemier, Hyung-Yong Kim & Gu Kong
Academic Unit of Surgery, School of Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow Royal Infirmary, Glasgow, UK
Nigel B. Jamieson
Department of Pathology, Asan Medical Center, College of Medicine, Ulsan University, Songpa-gu, Seoul, South Korea
Se Jin Jang & Hee Jin Lee
Science Writer, Garrett Park, MD, USA
Karine Jegalian
International Cancer Genome Consortium (ICGC)/ICGC Accelerating Research in Genomic Oncology (ARGO) Secretariat, Ontario Institute for Cancer Research, Toronto, ON, Canada
Jennifer L. Jennings
University of Ljubljana, Ljubljana, Slovenia
Lara Jerman
Department of Public Health Sciences, University of Chicago, Chicago, IL, USA
Yuan Ji
Research Institute, NorthShore University HealthSystem, Evanston, IL, USA
Yuan Ji
Department for Biomedical Research, University of Bern, Bern, Switzerland
Rory Johnson, Andrés Lanzós & Mark A. Rubin
Centre of Genomics and Policy, McGill University and Génome Québec Innovation Centre, Montreal, QC, Canada
Yann Joly, Bartha M. Knoppers, Mark Phillips & Adrian Thorogood
Carolina Center for Genome Sciences, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Corbin D. Jones
Hopp Children’s Cancer Center (KiTZ), Heidelberg, Germany
David T. W. Jones, Marcel Kool & Stefan M. Pfister
Pediatric Glioma Research Group, German Cancer Research Center (DKFZ), Heidelberg, Germany
David T. W. Jones
Cancer Research UK, London, UK
Nic Jones & David Scott
Indivumed GmbH, Hamburg, Germany
Hartmut Juhl
Genome Integration Data Center, Syntekabio, Inc, Daejeon, South Korea
Jongsun Jung
University Hospital Zurich, Zurich, Switzerland
Andre Kahles, Kjong-Van Lehmann & Gunnar Rätsch
Clinical Bioinformatics, Swiss Institute of Bioinformatics, Geneva, Switzerland
Abdullah Kahraman
Institute for Pathology and Molecular Pathology, University Hospital Zurich, Zurich, Switzerland
Abdullah Kahraman
Institute of Molecular Life Sciences, University of Zurich, Zurich, Switzerland
Abdullah Kahraman & Christian von Mering
MRC Human Genetics Unit, MRC IGMM, University of Edinburgh, Edinburgh, UK
Vera B. Kaiser & Colin A. Semple
Women’s Cancer Program at the Samuel Oschin Comprehensive Cancer Institute, Cedars-Sinai Medical Center, Los Angeles, CA, USA
Beth Karlan
Department of Biology, Bioinformatics Group, Division of Molecular Biology, Faculty of Science, University of Zagreb, Zagreb, Croatia
Rosa Karlić
Department for Internal Medicine II, University Hospital Schleswig-Holstein, Kiel, Germany
Dennis Karsch & Michael Kneba
Genetics and Molecular Pathology, SA Pathology, Adelaide, SA, Australia
Karin S. Kassahn
Department of Gastric Surgery, National Cancer Center Hospital, Tokyo, Japan
Hitoshi Katai
Department of Bioinformatics, Division of Cancer Genomics, National Cancer Center Research Institute, Tokyo, Japan
Mamoru Kato, Hirofumi Rokutan & Mihoko Saito-Adachi
A.A. Kharkevich Institute of Information Transmission Problems, Moscow, Russia
Marat D. Kazanov
Oncology and Immunology, Dmitry Rogachev National Research Center of Pediatric Hematology, Moscow, Russia
Marat D. Kazanov
Skolkovo Institute of Science and Technology, Moscow, Russia
Marat D. Kazanov
Department of Surgery, The George Washington University, School of Medicine and Health Science, Washington, DC, USA
Electron Kebebew
Endocrine Oncology Branch, Center for Cancer Research, National Cancer Institute, National Institutes of Health, Bethesda, MD, USA
Electron Kebebew
Melanoma Institute Australia, Macquarie University, Sydney, NSW, Australia
Richard F. Kefford
MIT Computer Science and Artificial Intelligence Laboratory, Massachusetts Institute of Technology, Cambridge, MA, USA
Manolis Kellis
Tissue Pathology and Diagnostic Oncology, Royal Prince Alfred Hospital, Sydney, NSW, Australia
James G. Kench & Richard A. Scolyer
Cholangiocarcinoma Screening and Care Program and Liver Fluke and Cholangiocarcinoma Research Centre, Faculty of Medicine, Khon Kaen University, Khon Kaen, Thailand
Narong Khuntikeo
Controlled Department and Institution, New York, NY, USA
Ekta Khurana
Englander Institute for Precision Medicine, Weill Cornell Medicine, New York, NY, USA
Ekta Khurana & Alexander Martinez-Fundichely
National Cancer Center, Gyeonggi, South Korea
Hark Kyun Kim
Department of Biochemistry, College of Medicine, Ewha Womans University, Seoul, South Korea
Hyung-Lae Kim
Health Sciences Department of Biomedical Informatics, University of California San Diego, La Jolla, CA, USA
Jihoon Kim
Research Core Center, National Cancer Centre Korea, Goyang-si, South Korea
Jong K. Kim
Department of Health Sciences and Technology, Sungkyunkwan University School of Medicine, Seoul, South Korea
Youngwook Kim
Samsung Genome Institute, Seoul, South Korea
Youngwook Kim
Breast Oncology Program, Dana-Farber/Brigham and Women’s Cancer Center, Boston, MA, USA
Tari A. King
Department of Surgery, Memorial Sloan Kettering Cancer Center, New York, NY, USA
Tari A. King & Samuel Singer
Division of Breast Surgery, Brigham and Women’s Hospital, Boston, MA, USA
Tari A. King
Integrative Bioinformatics Support Group, National Institute of Environmental Health Sciences (NIEHS), Durham, NC, USA
Leszek J. Klimczak
Department of Clinical Science, University of Bergen, Bergen, Norway
Stian Knappskog & Ola Myklebost
Center For Medical Innovation, Seoul National University Hospital, Seoul, South Korea
Youngil Koh
Department of Internal Medicine, Seoul National University Hospital, Seoul, South Korea
Youngil Koh & Sung-Soo Yoon
Institute of Computer Science, Polish Academy of Sciences, Warsawa, Poland
Jan Komorowski
Functional and Structural Genomics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Marcel Kool, Andrey Korshunov, Michael Koscher, Stefan M. Pfister & Qi Wang
Laboratory of Translational Genomics, Division of Cancer Epidemiology and Genetics, National Cancer Institute, , National Institutes of Health, Bethesda, MD, USA
Roelof Koster
Institute for Medical Informatics Statistics and Epidemiology, University of Leipzig, Leipzig, Germany
Markus Kreuz & Markus Loeffler
Morgan Welch Inflammatory Breast Cancer Research Program and Clinic, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Savitri Krishnamurthy
Department of Hematology and Oncology, Georg-Augusts-University of Göttingen, Göttingen, Germany
Dieter Kube & Lorenz H. P. Trümper
Institute of Cell Biology (Cancer Research), University of Duisburg-Essen, Essen, Germany
Ralf Küppers
King’s College London and Guy’s and St. Thomas’ NHS Foundation Trust, London, UK
Jesper Lagergren
Center for Epigenetics, Van Andel Research Institute, Grand Rapids, MI, USA
Peter W. Laird
The University of Queensland Centre for Clinical Research, Royal Brisbane and Women’s Hospital, Herston, QLD, Australia
Sunil R. Lakhani & Peter T. Simpson
Department of Pediatric Oncology and Hematology, University of Cologne, Cologne, Germany
Pablo Landgraf
University of Düsseldorf, Düsseldorf, Germany
Pablo Landgraf & Guido Reifenberger
Department of Pathology, Institut Jules Bordet, Brussels, Belgium
Denis Larsimont
Institute of Biomedicine, Sahlgrenska Academy at University of Gothenburg, Gothenburg, Sweden
Erik Larsson
Children’s Medical Research Institute, Sydney, NSW, Australia
Loretta M. S. Lau & Hilda A. Pickett
ILSbio, LLC Biobank, Chestertown, MD, USA
Xuan Le
Division of Genetics and Genomics, Boston Children’s Hospital, Harvard Medical School, Boston, MA, USA
Eunjung Alice Lee
Institute for Bioengineering and Biopharmaceutical Research (IBBR), Hanyang University, Seoul, South Korea
Jeong-Yeon Lee
Department of Statistics, University of California Santa Cruz, Santa Cruz, CA, USA
Juhee Lee
National Genotyping Center, Institute of Biomedical Sciences, Academia Sinica, Taipei, Taiwan
Ming Ta Michael Lee
Department of Vertebrate Genomics/Otto Warburg Laboratory Gene Regulation and Systems Biology of Cancer, Max Planck Institute for Molecular Genetics, Berlin, Germany
Hans Lehrach, Hans-Jörg Warnatz & Marie-Laure Yaspo
McGill University and Genome Quebec Innovation Centre, Montreal, QC, Canada
Louis Letourneau
biobyte solutions GmbH, Heidelberg, Germany
Ivica Letunic
Gynecologic Oncology, NYU Laura and Isaac Perlmutter Cancer Center, New York University, New York, NY, USA
Douglas A. Levine
Division of Oncology, Stem Cell Biology Section, Washington University School of Medicine, St. Louis, MO, USA
Tim Ley
Department of Systems Biology, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Han Liang
Harvard University, Cambridge, MA, USA
Ziao Lin
Urologic Oncology Branch, Center for Cancer Research, National Cancer Institute, National Institutes of Health, Bethesda, MD, USA
W. M. Linehan
University of Oslo, Oslo, Norway
Ole Christian Lingjærde & Torill Sauer
University of Toronto, Toronto, ON, Canada
Fei-Fei Fei Liu, Quaid D. Morris, Ruian Shi, Shankar Vembu & Fan Yang
Peking University, Beijing, China
Fenglin Liu, Fan Zhang, Liangtao Zheng & Xiuqing Zheng
School of Life Sciences, Peking University, Beijing, China
Fenglin Liu
Leidos Biomedical Research, Inc, McLean, VA, USA
Jia Liu
Hematology, Hospital Clinic, Institut d’Investigacions Biomèdiques August Pi i Sunyer (IDIBAPS), University of Barcelona, Barcelona, Spain
Armando Lopez-Guillermo
Second Military Medical University, Shanghai, China
Yong-Jie Lu & Hongwei Zhang
Chinese Cancer Genome Consortium, Shenzhen, China
Youyong Lu
Department of Medical Oncology, Beijing Hospital, Beijing, China
Youyong Lu
Laboratory of Molecular Oncology, Key Laboratory of Carcinogenesis and Translational Research (Ministry of Education), Peking University Cancer Hospital and Institute, Beijing, China
Youyong Lu & Rui Xing
School of Medicine/School of Mathematics and Statistics, University of St. Andrews, St, Andrews, Fife, UK
Andy G. Lynch
Institute for Systems Biology, Seattle, WA, USA
Lisa Lype, Sheila M. Reynolds & Ilya Shmulevich
Department of Biochemistry and Molecular Biology, Faculty of Medicine, University Institute of Oncology-IUOPA, Oviedo, Spain
Carlos López-Otín & Xose S. Puente
Institut Bergonié, Bordeaux, France
Gaetan MacGrogan
Cancer Unit, MRC University of Cambridge, Cambridge, UK
Shona MacRae
Department of Pathology and Laboratory Medicine, Center for Personalized Medicine, Children’s Hospital Los Angeles, Los Angeles, CA, USA
Dennis T. Maglinte
John Curtin School of Medical Research, Canberra, ACT, Australia
Graham J. Mann
MVZ Department of Oncology, PraxisClinic am Johannisplatz, Leipzig, Germany
Luisa Mantovani-Löffler
Department of Information Technology, Ghent University, Ghent, Belgium
Kathleen Marchal & Sergio Pulido-Tamayo
Department of Plant Biotechnology and Bioinformatics, Ghent University, Ghent, Belgium
Kathleen Marchal, Sergio Pulido-Tamayo & Lieven P. C. Verbeke
Institute for Genomic Medicine, Nationwide Children’s Hospital, Columbus, OH, USA
Elaine R. Mardis
Computational Biology Program, School of Medicine, Oregon Health and Science University, Portland, OR, USA
Adam A. Margolin & Adam J. Struck
Department of Surgery, Duke University, Durham, NC, USA
Jeffrey Marks
Institució Catalana de Recerca i Estudis Avançats (ICREA), Barcelona, Spain
Tomas Marques-Bonet, Jose I. Martin-Subero, Arcadi Navarro, David Torrents & Alfonso Valencia
Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Barcelona, Spain
Tomas Marques-Bonet
University of Glasgow, Glasgow, UK
Sancha Martin & Ke Yuan
Institut d’Investigacions Biomèdiques August Pi i Sunyer (IDIBAPS), Barcelona, Spain
Jose I. Martin-Subero
Division of Oncology, Washington University School of Medicine, St. Louis, MO, USA
R. Jay Mashl
Department of Surgery and Cancer, Imperial College, London, INY, UK
Erik Mayer
Applications Department, Oxford Nanopore Technologies, Oxford, UK
Simon Mayes & Daniel J. Turner
Department of Obstetrics, Gynecology and Reproductive Services, University of California San Francisco, San Francisco, CA, USA
Karen McCune & Karen Smith-McCune
Department of Biochemistry and Molecular Medicine, University California at Davis, Sacramento, CA, USA
John D. McPherson
STTARR Innovation Facility, Princess Margaret Cancer Centre, Toronto, ON, Canada
Alice Meng
Discipline of Surgery, Western Sydney University, Penrith, NSW, Australia
Neil D. Merrett
Yale School of Medicine, Yale University, New Haven, CT, USA
William Meyerson
Department of Genetics, Lineberger Comprehensive Cancer Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Piotr A. Mieczkowski, Joel S. Parker, Charles M. Perou, Donghui Tan, Umadevi Veluvolu & Matthew D. Wilkerson
Departments of Neurology and Neurosurgery, Henry Ford Hospital, Detroit, MI, USA
Tom Mikkelsen
Precision Oncology, OHSU Knight Cancer Institute, Oregon Health and Science University, Portland, OR, USA
Gordon B. Mills
Institute of Pathology, University Medical Center Hamburg-Eppendorf, Hamburg, Germany
Sarah Minner, Guido Sauter & Ronald Simon
Department of Health Sciences, Faculty of Medical Sciences, Kyushu University, Fukuoka, Japan
Shinichi Mizuno
Heidelberg Academy of Sciences and Humanities, Heidelberg, Germany
Fruzsina Molnár-Gábor
Department of Clinical Pathology, University of Melbourne, Melbourne, VIC, Australia
Carl Morrison, Karin A. Oien, Chawalit Pairojkul, Paul M. Waring & Marc J. van de Vijver
Department of Pathology, Roswell Park Cancer Institute, Buffalo, NY, USA
Carl Morrison
Department of Computer Science, University of Helsinki, Helsinki, Finland
Ville Mustonen
Institute of Biotechnology, University of Helsinki, Helsinki, Finland
Ville Mustonen
Organismal and Evolutionary Biology Research Programme, University of Helsinki, Helsinki, Finland
Ville Mustonen
Department of Obstetrics and Gynecology, Division of Gynecologic Oncology, Washington University School of Medicine, St. Louis, MO, USA
David Mutch
Penrose St. Francis Health Services, Colorado Springs, CO, USA
Jerome Myers
Institute of Pathology, Ulm University and University Hospital of Ulm, Ulm, Germany
Peter Möller
National Cancer Center, Tokyo, Japan
Hitoshi Nakagama
Genome Institute of Singapore, Singapore, Singapore
Tannistha Nandi & Patrick Tan
32Program in Computational Biology and Bioinformatics, Yale University, New Haven, CT, USA
Fabio C. P. Navarro
German Cancer Aid, Bonn, Germany
Gerd Nettekoven & Laura Planko
Programme in Cancer and Stem Cell Biology, Centre for Computational Biology, Duke-NUS Medical School, Singapore, Singapore
Alvin Wei Tian Ng
The Chinese University of Hong Kong, Shatin, NT, Hong Kong, China
Anthony Ng
Fourth Military Medical University, Shaanxi, China
Yongzhan Nie
The University of Cambridge School of Clinical Medicine, Cambridge, UK
Serena Nik-Zainal
St. Jude Children’s Research Hospital, Memphis, TN, USA
Paul A. Northcott
University Health Network, Princess Margaret Cancer Centre, Toronto, ON, Canada
Faiyaz Notta & Ming Tsao
Center for Biomolecular Science and Engineering, University of California Santa Cruz, Santa Cruz, CA, USA
Brian D. O’Connor
Department of Medicine, University of Chicago, Chicago, IL, USA
Peter O’Donnell
Department of Neurology, Mayo Clinic, Rochester, MN, USA
Brian Patrick O’Neill
Cambridge Oesophagogastric Centre, Cambridge University Hospitals NHS Foundation Trust, Cambridge, UK
J. Robert O’Neill
Department of Computer Science, Carleton College, Northfield, MN, USA
Layla Oesper
Institute of Cancer Sciences, College of Medical Veterinary and Life Sciences, University of Glasgow, Glasgow, UK
Karin A. Oien
Department of Epidemiology, University of Alabama at Birmingham, Birmingham, AL, USA
Akinyemi I. Ojesina
HudsonAlpha Institute for Biotechnology, Huntsville, AL, USA
Akinyemi I. Ojesina
O’Neal Comprehensive Cancer Center, University of Alabama at Birmingham, Birmingham, AL, USA
Akinyemi I. Ojesina
Department of Pathology, Keio University School of Medicine, Tokyo, Japan
Hidenori Ojima
Department of Hepatobiliary and Pancreatic Oncology, National Cancer Center Hospital, Tokyo, Japan
Takuji Okusaka
Sage Bionetworks, Seattle, WA, USA
Larsson Omberg
Lymphoma Genomic Translational Research Laboratory, National Cancer Centre, Singapore, Singapore
Choon Kiat Ong
Department of Clinical Pathology, Robert-Bosch-Hospital, Stuttgart, Germany
German Ott
Department of Cell and Systems Biology, University of Toronto, Toronto, ON, Canada
B. F. Francis Ouellette
Department of Biosciences and Nutrition, Karolinska Institutet, Stockholm, Sweden
Qiang Pan-Hammarström
Center for Liver Cancer, Research Institute and Hospital, National Cancer Center, Gyeonggi, South Korea
Joong-Won Park
Division of Hematology-Oncology, Samsung Medical Center, Sungkyunkwan University School of Medicine, Seoul, South Korea
Keunchil Park
Samsung Advanced Institute for Health Sciences and Technology, Sungkyunkwan University School of Medicine, Seoul, South Korea
Keunchil Park
Cheonan Industry-Academic Collaboration Foundation, Sangmyung University, Cheonan, South Korea
Kiejung Park
NYU Langone Medical Center, New York, NY, USA
Harvey Pass
Department of Hematology and Medical Oncology, Cleveland Clinic, Cleveland, OH, USA
Nathan A. Pennell
Department of Radiation Oncology, University of California San Francisco, San Francisco, CA, USA
Marc D. Perry
Department of Health Sciences Research, Mayo Clinic, Rochester, MN, USA
Gloria M. Petersen
Helen F. Graham Cancer Center at Christiana Care Health Systems, Newark, DE, USA
Nicholas Petrelli
Heidelberg University Hospital, Heidelberg, Germany
Stefan M. Pfister
CSRA Incorporated, Fairfax, VA, USA
Todd D. Pihl
Research Department of Pathology, University College London Cancer Institute, London, UK
Nischalan Pillay
Department of Research Oncology, Guy’s Hospital, King’s Health Partners AHSC, King’s College London School of Medicine, London, UK
Sarah Pinder
Faculty of Medicine and Health Sciences, Macquarie University, Sydney, NSW, Australia
Andreia V. Pinho
University Hospital of Minjoz, INSERM UMR 1098, Besançon, France
Xavier Pivot
Spanish National Cancer Research Centre, Madrid, Spain
Tirso Pons
Center of Digestive Diseases and Liver Transplantation, Fundeni Clinical Institute, Bucharest, Romania
Irinel Popescu
Cureline, Inc, South San Francisco, CA, USA
Olga Potapova
St. Luke’s Cancer Centre, Royal Surrey County Hospital NHS Foundation Trust, Guildford, UK
Shaun R. Preston
Cambridge Breast Unit, Addenbrooke’s Hospital, Cambridge University Hospital NHS Foundation Trust and NIHR Cambridge Biomedical Research Centre, Cambridge, UK
Elena Provenzano
East of Scotland Breast Service, Ninewells Hospital, Aberdeen, UK
Colin A. Purdie
Department of Genetics, Microbiology and Statistics, University of Barcelona, IRSJD, IBUB, Barcelona, Spain
Raquel Rabionet
Department of Obstetrics and Gynecology, Medical College of Wisconsin, Milwaukee, WI, USA
Janet S. Rader
Hematology and Medical Oncology, Winship Cancer Institute of Emory University, Atlanta, GA, USA
Suresh Ramalingam
Department of Computer Science, Princeton University, Princeton, NJ, USA
Benjamin J. Raphael & Matthew A. Reyna
Vanderbilt Ingram Cancer Center, Vanderbilt University, Nashville, TN, USA
W. Kimryn Rathmell
Ohio State University College of Medicine and Arthur G. James Comprehensive Cancer Center, Columbus, OH, USA
Matthew Ringel
Department of Surgery, Yokohama City University Graduate School of Medicine, Kanagawa, Japan
Yasushi Rino
Division of Chromatin Networks, German Cancer Research Center (DKFZ) and BioQuant, Heidelberg, Germany
Karsten Rippe
Research Computing Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Jeffrey Roach
School of Molecular Biosciences and Center for Reproductive Biology, Washington State University, Pullman, WA, USA
Steven A. Roberts
Finsen Laboratory and Biotech Research and Innovation Centre (BRIC), University of Copenhagen, Copenhagen, Denmark
F. Germán Rodríguez-González, Nikos Sidiropoulos & Joachim Weischenfeldt
Department of Laboratory Medicine and Pathobiology, University of Toronto, Toronto, ON, Canada
Michael H. A. Roehrl & Stefano Serra
Department of Pathology, Human Oncology and Pathogenesis Program, Memorial Sloan Kettering Cancer Center, New York, NY, USA
Michael H. A. Roehrl
University Hospital Giessen, Pediatric Hematology and Oncology, Giessen, Germany
Marius Rohde
Oncologie Sénologie, ICM Institut Régional du Cancer, Montpellier, France
Gilles Romieu
Institute of Clinical Molecular Biology, Christian-Albrechts-University, Kiel, Germany
Philip C. Rosenstiel & Markus B. Schilhabel
Institute of Pathology, University of Wuerzburg, Wuerzburg, Germany
Andreas Rosenwald
Department of Urology, North Bristol NHS Trust, Bristol, UK
Edward W. Rowe
SingHealth, Duke-NUS Institute of Precision Medicine, National Heart Centre Singapore, Singapore, Singapore
Steven G. Rozen, Patrick Tan & Bin Tean Teh
Department of Computer Science, University of Toronto, Toronto, ON, Canada
Yulia Rubanova, Jared T. Simpson & Jeffrey A. Wintersinger
Bern Center for Precision Medicine, University Hospital of Bern, University of Bern, Bern, Switzerland
Mark A. Rubin
Englander Institute for Precision Medicine, Weill Cornell Medicine and New York Presbyterian Hospital, New York, NY, USA
Mark A. Rubin
Meyer Cancer Center, Weill Cornell Medicine, New York, NY, USA
Mark A. Rubin
Pathology and Laboratory, Weill Cornell Medical College, New York, NY, USA
Mark A. Rubin
Vall d’Hebron Institute of Oncology: VHIO, Barcelona, Spain
Carlota Rubio-Perez
General and Hepatobiliary-Biliary Surgery, Pancreas Institute, University and Hospital Trust of Verona, Verona, Italy
Andrea Ruzzenente
National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bangalore, India
Radhakrishnan Sabarinathan
Indiana University, Bloomington, IN, USA
S. Cenk Sahinalp
Department of Pathology, GZA-ZNA Hospitals, Antwerp, Belgium
Roberto Salgado
Analytical Biological Services, Inc, Wilmington, DE, USA
Charles Saller
Sydney Medical School, University of Sydney, Sydney, NSW, Australia
Jaswinder S. Samra & Richard A. Scolyer
cBio Center, Dana-Farber Cancer Institute, Harvard Medical School, Boston, MA, USA
Chris Sander & Ciyue Shen
Department of Cell Biology, Harvard Medical School, Boston, MA, USA
Chris Sander & Ciyue Shen
Advanced Centre for Treatment Research and Education in Cancer, Tata Memorial Centre, Navi Mumbai, Maharashtra, India
Rajiv Sarin
School of Environmental and Life Sciences, Faculty of Science, The University of Newcastle, Ourimbah, NSW, Australia
Christopher J. Scarlett
Department of Dermatology, University Hospital of Essen, Essen, Germany
Dirk Schadendorf
Bioinformatics and Omics Data Analytics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Matthias Schlesner
Department of Urology, Charité Universitätsmedizin Berlin, Berlin, Germany
Thorsten Schlomm & Joachim Weischenfeldt
Martini-Clinic, Prostate Cancer Center, University Medical Center Hamburg-Eppendorf, Hamburg, Germany
Thorsten Schlomm
Department of General Internal Medicine, University of Kiel, Kiel, Germany
Stefan Schreiber
German Cancer Consortium (DKTK), Partner site Berlin, Berlin, Germany
Roland F. Schwarz
Cancer Research Institute, Beth Israel Deaconess Medical Center, Boston, MA, USA
Ralph Scully
University of Pittsburgh, Pittsburgh, PA, USA
Raja Seethala
Department of Ophthalmology and Ocular Genomics Institute, Massachusetts Eye and Ear, Harvard Medical School, Boston, MA, USA
Ayellet V. Segre
Center for Psychiatric Genetics, NorthShore University HealthSystem, Evanston, IL, USA
Subhajit Sengupta
Van Andel Research Institute, Grand Rapids, MI, USA
Hui Shen & Wanding Zhou
Laboratory of Molecular Medicine, Human Genome Center, Institute of Medical Science, University of Tokyo, Tokyo, Japan
Tatsuhiro Shibata, Hirokazu Taniguchi & Tomoko Urushidate
Japan Agency for Medical Research and Development, Tokyo, Japan
Kiyo Shimizu & Takashi Yugawa
Korea University, Seoul, South Korea
Seung Jun Shin & Stefan G. Stark
Murtha Cancer Center, Walter Reed National Military Medical Center, Bethesda, MD, USA
Craig Shriver
Human Genetics, University of Kiel, Kiel, Germany
Reiner Siebert
Department of Oncologic Pathology, Dana-Farber Cancer Institute, Harvard Medical School, Boston, MA, USA
Sabina Signoretti
Oregon Health and Science University, Portland, OR, USA
Jaclyn Smith
Center for RNA Interference and Noncoding RNA, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Anil K. Sood
Department of Experimental Therapeutics, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Anil K. Sood
Department of Gynecologic Oncology and Reproductive Medicine, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Anil K. Sood
University Hospitals Coventry and Warwickshire NHS Trust, Coventry, UK
Sharmila Sothi
Department of Radiation Oncology, Radboud University Nijmegen Medical Centre, Nijmegen, GA, The Netherlands
Paul N. Span
Institute for Genomics and Systems Biology, University of Chicago, Chicago, IL, USA
Jonathan Spring
Clinic for Hematology and Oncology, St.-Antonius-Hospital, Eschweiler, Germany
Peter Staib
Computational and Systems Biology Program, Memorial Sloan Kettering Cancer Center, New York, NY, USA
Stefan G. Stark
University of Iceland, Reykjavik, Iceland
Ólafur Andri Stefánsson
Division of Computational Genomics and Systems Genetics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Oliver Stegle
Dundee Cancer Centre, Ninewells Hospital, Dundee, UK
Alasdair Stenhouse & Alastair M. Thompson
Department for Internal Medicine III, University of Ulm and University Hospital of Ulm, Ulm, Germany
Stephan Stilgenbauer
Institut Curie, INSERM Unit 830, Paris, France
Henk G. Stunnenberg & Anne Vincent-Salomon
Department of Gastroenterology and Hepatology, Yokohama City University Graduate School of Medicine, Kanagawa, Japan
Akihiro Suzuki
Department of Laboratory Medicine, Radboud University Nijmegen Medical Centre, Nijmegen, GA, The Netherlands
Fred Sweep
Division of Cancer Genome Research, German Cancer Research Center (DKFZ), Heidelberg, Germany
Holger Sültmann
Department of General Surgery, Singapore General Hospital, Singapore, Singapore
Benita Kiat Tee Tan
Cancer Science Institute of Singapore, National University of Singapore, Singapore, Singapore
Patrick Tan & Bin Tean Teh
Department of Medical and Clinical Genetics, Genome-Scale Biology Research Program, University of Helsinki, Helsinki, Finland
Tomas J. Tanskanen
East Anglian Medical Genetics Service, Cambridge University Hospitals NHS Foundation Trust, Cambridge, UK
Patrick Tarpey
Irving Institute for Cancer Dynamics, Columbia University, New York, NY, USA
Simon Tavaré
Institute of Molecular and Cell Biology, Singapore, Singapore
Bin Tean Teh
Laboratory of Cancer Epigenome, Division of Medical Science, National Cancer Centre Singapore, Singapore, Singapore
Bin Tean Teh
Universite Lyon, INCa-Synergie, Centre Léon Bérard, Lyon, France
Gilles Thomas
Department of Urology, Mayo Clinic, Rochester, MN, USA
R. Houston Thompson
Royal National Orthopaedic Hospital - Stanmore, Stanmore, Middlesex, UK
Roberto Tirabosco
Department of Biochemistry, Genetics and Immunology, University of Vigo, Vigo, Spain
Marta Tojo
Giovanni Paolo II / I.R.C.C.S. Cancer Institute, Bari, BA, Italy
Stefania Tommasi
Neuroblastoma Genomics, German Cancer Research Center (DKFZ), Heidelberg, Germany
Umut H. Toprak
Fondazione Policlinico Universitario Gemelli IRCCS, Rome, Italy, Rome, Italy
Giampaolo Tortora
University of Verona, Verona, Italy
Giampaolo Tortora
Centre National de Génotypage, CEA - Institute de Génomique, Evry, France
Jörg Tost
CAPHRI Research School, Maastricht University, Maastricht, ER, The Netherlands
David Townend
Department of Biopathology, Centre Léon Bérard, Lyon, France
Isabelle Treilleux
Université Claude Bernard Lyon 1, Villeurbanne, France
Isabelle Treilleux
Core Research for Evolutional Science and Technology (CREST), JST, Tokyo, Japan
Tatsuhiko Tsunoda
Department of Biological Sciences, Laboratory for Medical Science Mathematics, Graduate School of Science, University of Tokyo, Yokohama, Japan
Tatsuhiko Tsunoda
Department of Medical Science Mathematics, Medical Research Institute, Tokyo Medical and Dental University (TMDU), Tokyo, Japan
Tatsuhiko Tsunoda
Cancer Ageing and Somatic Mutation Programme, Wellcome Sanger Institute, Hinxton, UK
Jose M. C. Tubio
University Hospitals Birmingham NHS Foundation Trust, Birmingham, UK
Olga Tucker
Centre for Cancer Research and Cell Biology, Queen’s University, Belfast, UK
Richard Turkington
Breast Medical Oncology, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Naoto T. Ueno
Department of Surgery, Johns Hopkins University School of Medicine, Baltimore, MD, USA
Christopher Umbricht
Department of Oncology-Pathology, Science for Life Laboratory, Karolinska Institute, Stockholm, Sweden
Husen M. Umer
School of Cancer Sciences, Faculty of Medicine, University of Southampton, Southampton, UK
Timothy J. Underwood
Department of Gene Technology, Tallinn University of Technology, Tallinn, Estonia
Liis Uusküla-Reimand
Genetics and Genome Biology Program, SickKids Research Institute, The Hospital for Sick Children, Toronto, ON, Canada
Liis Uusküla-Reimand
Departments of Neurosurgery and Hematology and Medical Oncology, Winship Cancer Institute and School of Medicine, Emory University, Atlanta, GA, USA
Erwin G. Van Meir
Department of Clinical and Molecular Medicine, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology, Trondheim, Norway
Miguel Vazquez
Argmix Consulting, North Vancouver, BC, Canada
Shankar Vembu
Department of Information Technology, Ghent University, Interuniversitair Micro-Electronica Centrum (IMEC), Ghent, Belgium
Lieven P. C. Verbeke
Nuffield Department of Surgical Sciences, John Radcliffe Hospital, University of Oxford, Oxford, UK
Clare Verrill
Institute of Mathematics and Computer Science, University of Latvia, Riga, LV, Latvia
Juris Viksna
Discipline of Pathology, Sydney Medical School, University of Sydney, Sydney, NSW, Australia
Ricardo E. Vilain
Department of Applied Mathematics and Theoretical Physics, Centre for Mathematical Sciences, University of Cambridge, Cambridge, UK
Ignacio Vázquez-García
Department of Epidemiology and Biostatistics, Memorial Sloan Kettering Cancer Center, New York, NY, USA
Ignacio Vázquez-García & Venkata D. Yellapantula
Department of Statistics, Columbia University, New York, NY, USA
Ignacio Vázquez-García
Department of Immunology, Genetics and Pathology, Science for Life Laboratory, Uppsala University, Uppsala, Sweden
Claes Wadelius
School of Electronic and Information Engineering, Xi’an Jiaotong University, Xi’an, China
Jiayin Wang & Kai Ye
Department of Histopathology, Cambridge University Hospitals NHS Foundation Trust, Cambridge, UK
Anne Y. Warren
Oxford NIHR Biomedical Research Centre, University of Oxford, Oxford, UK
David C. Wedge
Georgia Regents University Cancer Center, Augusta, GA, USA
Paul Weinberger
Wythenshawe Hospital, Manchester, UK
Ian Welch
Department of Genetics, Washington University School of Medicine, St.Louis, MO, USA
Michael C. Wendl
Department of Biological Oceanography, Leibniz Institute of Baltic Sea Research, Rostock, Germany
Johannes Werner
Wellcome Centre for Human Genetics, University of Oxford, Oxford, UK
Justin P. Whalley
Department of Molecular and Human Genetics, Baylor College of Medicine, Houston, TX, USA
David A. Wheeler
Thoracic Oncology Laboratory, Mayo Clinic, Rochester, MN, USA
Dennis Wigle
Institute for Genomic Medicine, Nationwide Children’s Hospital, Columbus, OH, USA
Richard K. Wilson
Department of Obstetrics and Gynecology, Division of Gynecologic Oncology, Mayo Clinic, Rochester, MN, USA
Boris Winterhoff
International Institute for Molecular Oncology, Poznań, Poland
Maciej Wiznerowicz
Poznan University of Medical Sciences, Poznań, Poland
Maciej Wiznerowicz
Genomics and Proteomics Core Facility High Throughput Sequencing Unit, German Cancer Research Center (DKFZ), Heidelberg, Germany
Stephan Wolf
NCCS-VARI Translational Research Laboratory, National Cancer Centre Singapore, Singapore, Singapore
Bernice H. Wong
Edison Family Center for Genome Sciences and Systems Biology, Washington University, St. Louis, MO, USA
Winghing Wong
MRC-University of Glasgow Centre for Virus Research, Glasgow, UK
Derek W. Wright
Department of Medical Informatics and Clinical Epidemiology, Division of Bioinformatics and Computational Biology, OHSU Knight Cancer Institute, Oregon Health and Science University, Portland, OR, USA
Guanming Wu
School of Electronic Information and Communications, Huazhong University of Science and Technology, Wuhan, China
Tian Xia
Department of Applied Mathematics and Statistics, Johns Hopkins University, Baltimore, MD, USA
Yanxun Xu
Department of Cancer Genome Informatics, Graduate School of Medicine, Osaka University, Osaka, Japan
Shinichi Yachida
Institute of Computer Science, Heidelberg University, Heidelberg, Germany
Sergei Yakneen
School of Mathematics and Statistics, University of Sydney, Sydney, NSW, Australia
Jean Y. Yang
Ben May Department for Cancer Research, University of Chicago, Chicago, IL, USA
Lixing Yang
Department of Human Genetics, University of Chicago, Chicago, IL, USA
Lixing Yang
Tri-Institutional PhD Program in Computational Biology and Medicine, Weill Cornell Medicine, New York, NY, USA
Xiaotong Yao
The First Affiliated Hospital, Xi’an Jiaotong University, Xi’an, China
Kai Ye
Department of Medicine and Therapeutics, The Chinese University of Hong Kong, Shatin, NT, Hong Kong, China
Jun Yu
Department of Biostatistics, The University of Texas MD Anderson Cancer Center, Houston, TX, USA
Kaixian Yu & Hongtu Zhu
Duke-NUS Medical School, Singapore, Singapore
Willie Yu
Department of Surgery, Ruijin Hospital, Shanghai Jiaotong University School of Medicine, Shanghai, China
Yingyan Yu
School of Computing Science, University of Glasgow, Glasgow, UK
Ke Yuan
Division of Orthopaedic Surgery, Oslo University Hospital, Oslo, Norway
Olga Zaikova
Eastern Clinical School, Monash University, Melbourne, VIC, Australia
Nikolajs Zeps
Epworth HealthCare, Richmond, VIC, Australia
Nikolajs Zeps
Department of Biostatistics and Computational Biology, Dana-Farber Cancer Institute and Harvard Medical School, Boston, MA, USA
Cheng-Zhong Zhang
Department of Biomedical Informatics, College of Medicine, The Ohio State University, Columbus, OH, USA
Yan Zhang
The Ohio State University Comprehensive Cancer Center (OSUCCC – James), Columbus, OH, USA
Yan Zhang
The University of Texas School of Biomedical Informatics (SBMI) at Houston, Houston, TX, USA
Zhongming Zhao
Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
Hongtu Zhu
Department of Biochemistry and Molecular Genetics, Feinberg School of Medicine, Northwestern University, Chicago, IL, USA
Lihua Zou
Faculty of Medicine and Health, University of Sydney, Sydney, NSW, Australia
Anna deFazio
Department of Pathology, Erasmus Medical Center Rotterdam, Rotterdam, GD, The Netherlands
Carolien H. M. van Deurzen
Division of Molecular Carcinogenesis, The Netherlands Cancer Institute, Amsterdam, CX, The Netherlands
L. van’t Veer
Institute of Molecular Life Sciences and Swiss Institute of Bioinformatics, University of Zurich, Zurich, Switzerland
Christian von Mering

Authors

Vinayak Bhandari
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Constance H. Li
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Robert G. Bristow
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Paul C. Boutros
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Consortia

PCAWG Consortium

Lauri A. Aaltonen
, Federico Abascal
, Adam Abeshouse
, Hiroyuki Aburatani
, David J. Adams
, Nishant Agrawal
, Keun Soo Ahn
, Sung-Min Ahn
, Hiroshi Aikata
, Rehan Akbani
, Kadir C. Akdemir
, Hikmat Al-Ahmadie
, Sultan T. Al-Sedairy
, Fatima Al-Shahrour
, Malik Alawi
, Monique Albert
, Kenneth Aldape
, Ludmil B. Alexandrov
, Adrian Ally
, Kathryn Alsop
, Eva G. Alvarez
, Fernanda Amary
, Samirkumar B. Amin
, Brice Aminou
, Ole Ammerpohl
, Matthew J. Anderson
, Yeng Ang
, Davide Antonello
, Pavana Anur
, Samuel Aparicio
, Elizabeth L. Appelbaum
, Yasuhito Arai
, Axel Aretz
, Koji Arihiro
, Shun-ichi Ariizumi
, Joshua Armenia
, Laurent Arnould
, Sylvia Asa
, Yassen Assenov
, Gurnit Atwal
, Sietse Aukema
, J. Todd Auman
, Miriam R. R. Aure
, Philip Awadalla
, Marta Aymerich
, Gary D. Bader
, Adrian Baez-Ortega
, Matthew H. Bailey
, Peter J. Bailey
, Miruna Balasundaram
, Saianand Balu
, Pratiti Bandopadhayay
, Rosamonde E. Banks
, Stefano Barbi
, Andrew P. Barbour
, Jonathan Barenboim
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, Hugh Barr
, Elisabet Barrera
, John Bartlett
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, Claudio Bassi
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, Stephen B. Baylin
, Wojciech Bazant
, Duncan Beardsmore
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, Benedikt Brors
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, Alicia L. Bruzos
, Alex Buchanan
, Ivo Buchhalter
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, Niall J. Byrne
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, Samantha J. Caesar-Johnson
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, Declan Cahill
, Claudia Calabrese
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, Fabien Calvo
, Niedzica Camacho
, Peter J. Campbell
, Elias Campo
, Cinzia Cantù
, Shaolong Cao
, Thomas E. Carey
, Joana Carlevaro-Fita
, Rebecca Carlsen
, Ivana Cataldo
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, Jonathan Cebon
, Robert Cerfolio
, Dianne E. Chadwick
, Dimple Chakravarty
, Don Chalmers
, Calvin Wing Yiu Chan
, Kin Chan
, Michelle Chan-Seng-Yue
, Vishal S. Chandan
, David K. Chang
, Stephen J. Chanock
, Lorraine A. Chantrill
, Aurélien Chateigner
, Nilanjan Chatterjee
, Kazuaki Chayama
, Hsiao-Wei Chen
, Jieming Chen
, Ken Chen
, Yiwen Chen
, Zhaohong Chen
, Andrew D. Cherniack
, Jeremy Chien
, Yoke-Eng Chiew
, Suet-Feung Chin
, Juok Cho
, Sunghoon Cho
, Jung Kyoon Choi
, Wan Choi
, Christine Chomienne
, Zechen Chong
, Su Pin Choo
, Angela Chou
, Angelika N. Christ
, Elizabeth L. Christie
, Eric Chuah
, Carrie Cibulskis
, Kristian Cibulskis
, Sara Cingarlini
, Peter Clapham
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, Nicole Cloonan
, Marek Cmero
, Colin C. Collins
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, Brian Craft
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, Chad J. Creighton
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, Erin Curley
, Ioana Cutcutache
, Karolina Czajka
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, Rebecca A. Dagg
, Ludmila Danilova
, Maria Vittoria Davi
, Natalie R. Davidson
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, Kevin J. Dawson
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, Deniz Demircioğlu
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, Stefan C. Dentro
, Nikita Desai
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, Anthony DiBiase
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, Adel El-Naggar
, Matthew Eldridge
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, Serap Erkek
, Georgia Escaramis
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, Dariush Etemadmoghadam
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, Robert S. Fulton
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, Anja Füllgrabe
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, Levi Garraway
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, Nicola Sperandio
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Contributions

Bioinformatics and statistical analysis: V.B. Data processing: V.B., C.H.L. Data visualization: V.B. Supervised research: R.G.B., P.C.B. Initiated the project: V.B., P.C.B. Wrote the first draft of the manuscript: V.B. Approved the manuscript: all authors.

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Correspondence to Robert G. Bristow or Paul C. Boutros.

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Bhandari, V., Li, C.H., Bristow, R.G. et al. Divergent mutational processes distinguish hypoxic and normoxic tumours. Nat Commun 11, 737 (2020). https://doi.org/10.1038/s41467-019-14052-x

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Received: 02 May 2019
Accepted: 29 November 2019
Published: 05 February 2020
DOI: https://doi.org/10.1038/s41467-019-14052-x

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