From c80dd80dd7bcddad031c6c972bb6403d554e90fc Mon Sep 17 00:00:00 2001 From: John Huddleston Date: Tue, 13 Aug 2024 16:25:57 -0700 Subject: [PATCH 1/8] Add initial within-clade cluster workflow/text Adds initial results text from a prototype analysis as a placeholder and adds rules to the early SC2 workflow to automate the prototype analysis, generating accuracy files for t-SNE clusters of 21J (Delta) sequences. Results text will definitely change as we rerun this analysis. Closes #110 --- manuscript/cartography.tex | 11 ++ sars-cov-2-nextstrain/Snakefile | 214 ++++++++++++++++++++++++++++++++ 2 files changed, 225 insertions(+) diff --git a/manuscript/cartography.tex b/manuscript/cartography.tex index 84d311ce..ef72abbe 100644 --- a/manuscript/cartography.tex +++ b/manuscript/cartography.tex @@ -436,6 +436,17 @@ \subsection{SARS-CoV-2 clusters recapitulate broad genetic groups corresponding With the exception of t-SNE's performance, these results replicate the patterns we observed with early SARS-CoV-2 data where clusters from embeddings more effectively represented broader genetic diversity than the finer resolution diversity denoted by Pango lineages. Unlike the Pango lineages in the early SARS-CoV-2 data, the lineages from the later data exhibited fewer pairwise genetic distances between samples in each lineage than samples in Nextstrain clades or any embedding cluster (Supplementary Fig.~S\ref{S_Fig_sarscov2_within_between_group_distances}). +To understand whether t-SNE clusters could capture Pango-resolution genetic groups within a single Nextstrain clade, we evenly sampled approximately 2,000 sequences from a dominant Nextstrain clade with many Pango lineages, 21J (Delta), and identified clusters from a t-SNE embedding of those data. +Within the 1,992 sequences from 21J (Delta), we found 37 Pango lineages after collapsing lineages with fewer than 10 sequences into their parent lineages. +We found 44 t-SNE clusters representing 1,466 sequences with 526 sequences (26\%) assigned to the unclustered ``-1'' label. +The VI distance between all clusters (including the ``-1'' group) and the Pango lineages was 0.16. +This distance was consistent with the distance of 0.14 between t-SNE clusters and Pango lineages from both the full early and late SARS-CoV-2 datasets. +The VI distance between clusters without the -1 group and Pango lineages was 0.07, confirming that half of the distance between t-SNE clusters and Pango lineages above came from unclustered sequences. +Of the 35 Pango lineages with a t-SNE cluster, 21 lineages (60\%) had a single corresponding t-SNE cluster and 12 lineages (34\%) had 2 or 3 t-SNE clusters. +The remaining two lineages, AY.4 and B.1.617.2, had 5 and 10 t-SNE clusters, respectively. +Of the 44 t-SNE clusters, 29 clusters (66\%) had a single corresponding Pango lineage, 14 (32\%) mapped to two Pango lineages, and one cluster (43) mapped to 6 Pango lineages. +These results suggest that clusters from t-SNE embeddings can capture Pango-resolution genetic groups when analyzing sequences within a specific Nextstrain clade. + \subsection{Distance-based embeddings reflect SARS-CoV-2 recombination events} Finally, we tested the ability of sequence embeddings to place known recombinant lineages of SARS-CoV-2 between their parental lineages in Euclidean space. diff --git a/sars-cov-2-nextstrain/Snakefile b/sars-cov-2-nextstrain/Snakefile index 3edc8a50..b8f73170 100644 --- a/sars-cov-2-nextstrain/Snakefile +++ b/sars-cov-2-nextstrain/Snakefile @@ -17,6 +17,9 @@ rule sarscov2: "auspice/cartography_ncov-2020-2022.json", expand("sars-cov-2-nextstrain/results/mutation_table_for_{clade_membership}.csv", clade_membership=SARS_COV_2_CLADE_MEMBERSHIPS), "sars-cov-2-nextstrain/results/monophyletic_clusters.csv", + "sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_t-sne_for_Nextclade_pango_collapsed.csv", + "sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_without_unclustered_t-sne_for_Nextclade_pango_collapsed.csv", + "sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts.tsv", rule sarscov2_download_filtered_metadata: output: @@ -960,3 +963,214 @@ rule sarscov2_get_authors_for_accessions: | csvtk rename -l -t -f "Isolate Lineage" -n "Virus Name" \ | csvtk mutate2 -l -t -n "dataset" -e "'sars-cov-2-2020-2022'" > {output.authors} """ + +rule sarscov2_subsample_single_clade: + input: + metadata="sars-cov-2-nextstrain/data/filtered_metadata.tsv.zst", + outliers="sars-cov-2-nextstrain/config/exclude.txt", + output: + strains="sars-cov-2-nextstrain/data/single_clade/strains.txt", + params: + random_seed=RANDOM_SEED, + single_clade="21J (Delta)", + conda: "../cartography.yml" + shell: + """ + augur filter \ + --metadata {input.metadata} \ + --exclude {input.outliers} \ + --min-date 2020-01-01 \ + --max-date 2022-01-01 \ + --query "Nextstrain_clade == '{params.single_clade}'" \ + --group-by region year month \ + --subsample-max-sequences 2000 \ + --subsample-seed {params.random_seed} \ + --output-strains {output.strains} + """ + +rule sarscov2_extract_subsampled_data_for_single_clade: + input: + metadata="sars-cov-2-nextstrain/data/filtered_metadata.tsv.zst", + alignment="sars-cov-2-nextstrain/data/all_aligned.fasta.zst", + strains="sars-cov-2-nextstrain/data/single_clade/strains.txt", + output: + metadata="sars-cov-2-nextstrain/data/single_clade/metadata.tsv", + alignment="sars-cov-2-nextstrain/data/single_clade/aligned.fasta", + conda: "../cartography.yml" + shell: + """ + augur filter \ + --metadata {input.metadata} \ + --sequences {input.alignment} \ + --exclude-all \ + --include {input.strains} \ + --output-metadata {output.metadata} \ + --output-sequences {output.alignment} + """ + +rule sarscov2_collapse_pango_lineages_for_single_clade: + input: + metadata="sars-cov-2-nextstrain/data/single_clade/metadata.tsv", + output: + metadata="sars-cov-2-nextstrain/data/single_clade/metadata_Pango_collapsed.tsv", + conda: "../cartography.yml" + log: + "logs/sars-cov-2-nextstrain/collapse_pango_lineages_for_single_clade.txt", + params: + min_samples=10, + pango_column="Nextclade_pango", + new_column="Nextclade_pango_collapsed", + shell: + """ + python3 notebooks/scripts/collapse_pango_lineages.py \ + --metadata {input.metadata} \ + --min-samples {params.min_samples} \ + --pango-column {params.pango_column} \ + --new-column {params.new_column} \ + --output {output.metadata} 2>&1 | tee {log} + """ + +rule sarscov2_create_distance_matrix_for_single_clade: + input: + alignment="sars-cov-2-nextstrain/data/single_clade/aligned.fasta", + output: + output="sars-cov-2-nextstrain/results/single_clade/distance_matrix.csv", + conda: "../cartography.yml" + shell: + """ + pathogen-distance \ + --alignment {input.alignment} \ + --indel-distance \ + --output {output.output} + """ + +rule sarscov2_embed_tsne_for_single_clade: + input: + alignment="sars-cov-2-nextstrain/data/single_clade/aligned.fasta", + distance_matrix="sars-cov-2-nextstrain/results/single_clade/distance_matrix.csv", + parameters="simulations/coronavirus-like/moderate-recombination-rate/t-sne_parameters.csv", + output: + dataframe="sars-cov-2-nextstrain/results/single_clade/embed_t-sne.csv", + params: + random_seed=RANDOM_SEED, + conda: "../cartography.yml" + shell: + """ + pathogen-embed \ + --alignment {input.alignment} \ + --distance-matrix {input.distance_matrix} \ + --embedding-parameters {input.parameters} \ + --random-seed {params.random_seed} \ + --output-dataframe {output.dataframe} \ + t-sne \ + --pca-encoding simplex + """ + +rule sarscov2_cluster_for_single_clade: + input: + embedding="sars-cov-2-nextstrain/results/single_clade/embed_t-sne.csv", + parameters="sars-cov-2-nextstrain/results/optimal_cluster_accuracy_and_parameters_for_Nextclade_pango_collapsed.csv", + output: + clustered_embedding="sars-cov-2-nextstrain/results/single_clade/cluster_embed_t-sne_for_Nextclade_pango_collapsed.csv", + params: + min_size=CLUSTER_MIN_SIZE, + min_samples=CLUSTER_MIN_SAMPLES, + conda: "../cartography.yml" + shell: + """ + pathogen-cluster \ + --embedding {input.embedding} \ + --min-size {params.min_size} \ + --min-samples {params.min_samples} \ + --distance-threshold "$(csvtk filter2 -f '$method=="t-sne"' {input.parameters} | csvtk cut -f distance_threshold | csvtk del-header)" \ + --label-attribute "t-sne_label_for_Nextclade_pango_collapsed" \ + --output-dataframe {output.clustered_embedding} + """ + +rule sarscov2_cluster_accuracy_for_single_clade: + input: + metadata="sars-cov-2-nextstrain/data/single_clade/metadata_Pango_collapsed.tsv", + clusters="sars-cov-2-nextstrain/results/single_clade/cluster_embed_t-sne_for_Nextclade_pango_collapsed.csv", + output: + dataframe="sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_t-sne_for_Nextclade_pango_collapsed.csv", + params: + ignored_clusters="unassigned", + conda: "../cartography.yml" + shell: + """ + python3 notebooks/scripts/metadata_HDBSCAN.py \ + --method "t-sne" \ + --true-clusters {input.metadata} \ + --true-clusters-column "Nextclade_pango_collapsed" \ + --predicted-clusters {input.clusters} \ + --predicted-clusters-column "t-sne_label_for_Nextclade_pango_collapsed" \ + --ignored-clusters {params.ignored_clusters:q} \ + --output {output.dataframe} + """ + +rule sarscov2_cluster_accuracy_without_unclustered_for_single_clade: + input: + metadata="sars-cov-2-nextstrain/data/single_clade/metadata_Pango_collapsed.tsv", + clusters="sars-cov-2-nextstrain/results/single_clade/cluster_embed_t-sne_for_Nextclade_pango_collapsed.csv", + output: + dataframe="sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_without_unclustered_t-sne_for_Nextclade_pango_collapsed.csv", + params: + ignored_clusters=["unassigned", "-1"], + conda: "../cartography.yml" + shell: + """ + python3 notebooks/scripts/metadata_HDBSCAN.py \ + --method "t-sne" \ + --true-clusters {input.metadata} \ + --true-clusters-column "Nextclade_pango_collapsed" \ + --predicted-clusters {input.clusters} \ + --predicted-clusters-column "t-sne_label_for_Nextclade_pango_collapsed" \ + --ignored-clusters {params.ignored_clusters:q} \ + --output {output.dataframe} + """ + +rule sarscov2_select_lineages_for_single_clade: + input: + metadata="sars-cov-2-nextstrain/data/single_clade/metadata_Pango_collapsed.tsv", + output: + lineages="sars-cov-2-nextstrain/results/single_clade/lineages.tsv", + conda: "../cartography.yml" + shell: + """ + tsv-select -H -f strain,Nextclade_pango_collapsed {input.metadata} > {output.lineages} + """ + +rule sarscov2_select_clusters_for_single_clade: + input: + clusters="sars-cov-2-nextstrain/results/single_clade/cluster_embed_t-sne_for_Nextclade_pango_collapsed.csv", + output: + clusters="sars-cov-2-nextstrain/results/single_clade/clusters.tsv", + conda: "../cartography.yml" + shell: + """ + tsv-select -d "," -H -f 'strain','t\-sne_label_for_Nextclade_pango_collapsed' {input.clusters} \ + | csv2tsv > {output.clusters} + """ + +rule sarscov2_merge_lineages_and_clusters_for_single_clade: + input: + lineages="sars-cov-2-nextstrain/results/single_clade/lineages.tsv", + clusters="sars-cov-2-nextstrain/results/single_clade/clusters.tsv", + output: + table="sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters.tsv", + conda: "../cartography.yml" + shell: + """ + tsv-join -H -f {input.lineages} -k strain -a Nextclade_pango_collapsed {input.clusters} > {output.table} + """ + +rule sarscov2_summarize_lineages_and_clusters_for_single_clade: + input: + table="sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters.tsv", + output: + counts="sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts.tsv", + conda: "../cartography.yml" + shell: + """ + tsv-summarize -H -g 'Nextclade_pango_collapsed','t\-sne_label_for_Nextclade_pango_collapsed' --count {input.table} > {output.counts} + """ From a4ed8cc957f9d8c050bfe5f9c828447bb2a4cd27 Mon Sep 17 00:00:00 2001 From: John Huddleston Date: Wed, 14 Aug 2024 13:35:23 -0700 Subject: [PATCH 2/8] Add results for single clade cluster accuracy Adds results tables and figures for single clade cluster analysis including rules to filter the lineage/cluster counts to exclude unclustered sequences and plot the lineage/cluster counts for the unfiltered table. --- ...gle_clade_lineages_and_clusters_counts.png | Bin 0 -> 79680 bytes sars-cov-2-nextstrain/Snakefile | 31 + ...cy_t-sne_for_Nextclade_pango_collapsed.csv | 2 + ...ed_t-sne_for_Nextclade_pango_collapsed.csv | 2 + .../single_clade/lineages_and_clusters.tsv | 1993 +++++++++++++++++ .../lineages_and_clusters_counts.tsv | 70 + ...nd_clusters_counts_without_unclustered.tsv | 53 + ...single-clade-lineage-and-cluster-counts.py | 28 + 8 files changed, 2179 insertions(+) create mode 100644 manuscript/figures/sarscov2_single_clade_lineages_and_clusters_counts.png create mode 100644 sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_t-sne_for_Nextclade_pango_collapsed.csv create mode 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counts="sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts_without_unclustered.tsv", + conda: "../cartography.yml" + shell: + """ + tsv-filter -H --str-ne "t\-sne_label_for_Nextclade_pango_collapsed:-1" {input.counts} > {output.counts} + """ + +rule sarscov2_plot_lineages_and_clusters_for_single_clade: + input: + counts="sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts.tsv", + output: + counts_plot="manuscript/figures/sarscov2_single_clade_lineages_and_clusters_counts.png", + conda: "../cartography.yml" + params: + lineage_column="Nextclade_pango_collapsed", + cluster_column="t-sne_label_for_Nextclade_pango_collapsed", + shell: + """ + python3 sars-cov-2-nextstrain/scripts/plot-single-clade-lineage-and-cluster-counts.py \ + --counts {input.counts} \ + --lineage-column {params.lineage_column:q} \ + --cluster-column {params.cluster_column:q} \ + --output {output.counts_plot} + """ diff --git a/sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_t-sne_for_Nextclade_pango_collapsed.csv b/sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_t-sne_for_Nextclade_pango_collapsed.csv new file mode 100644 index 00000000..71f3d76e --- /dev/null +++ b/sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_t-sne_for_Nextclade_pango_collapsed.csv @@ -0,0 +1,2 @@ +method,predicted_clusters_column,normalized_vi,n_predicted_clusters,n_predicted_cluster_samples,n_ignored_predicted_clusters,n_true_cluster_samples,n_ignored_true_clusters,n_vi_cluster_samples +t-sne,t-sne_label_for_Nextclade_pango_collapsed,0.17,28,1992,0,1992,0,1992 diff --git a/sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_without_unclustered_t-sne_for_Nextclade_pango_collapsed.csv b/sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_without_unclustered_t-sne_for_Nextclade_pango_collapsed.csv new file mode 100644 index 00000000..bc373a63 --- /dev/null +++ b/sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_without_unclustered_t-sne_for_Nextclade_pango_collapsed.csv @@ -0,0 +1,2 @@ +method,predicted_clusters_column,normalized_vi,n_predicted_clusters,n_predicted_cluster_samples,n_ignored_predicted_clusters,n_true_cluster_samples,n_ignored_true_clusters,n_vi_cluster_samples +t-sne,t-sne_label_for_Nextclade_pango_collapsed,0.13,28,1806,186,1992,0,1806 diff --git a/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters.tsv b/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters.tsv new file mode 100644 index 00000000..cede273b --- /dev/null +++ b/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters.tsv @@ -0,0 +1,1993 @@ +strain t-sne_label_for_Nextclade_pango_collapsed Nextclade_pango_collapsed +BRA/SP-NVBS6920GENOV828076619274/2021 1 AY.99.2 +USA/TX-CDC-LC0056011/2021 27 B.1.617.2 +England/LOND-YYBNF59/2022 27 AY.4.2 +LC752099 16 AY.29 +BHR/400215521_S32_L001/2021 27 B.1.617.2 +OY530201 27 AY.4 +England/NORW-13DACBA/2021 27 AY.4 +Northern_Ireland/NIRE-0050d0/2021 27 AY.4 +USA/MN-CDC-ASC210453623/2021 18 AY.3 +BHR/21342818722_S65_L001/2021 15 AY.3 +CHL/AR-UFRO-0421/2021 11 AY.101 +SouthAfrica/NHLS-UCT-GS-F132/2021 26 AY.45 +Switzerland/SO-ETHZ-33760694/2021 21 AY.43 +England/NORT-YYBGBS/2021 27 AY.4 +OV115124 27 AY.33 +NZL/21MV0277/2021 27 AY.4 +NZL/21CV0251/2021 9 AY.39.1.1 +USA/NC-CDC-STM-000069068/2021 27 B.1.617.2 +Japan/SZ-NIG-Y211710/2021 16 AY.29 +IND/hospital_2_130/2021 27 B.1.617.2 +OY628730 27 AY.4 +NZL/21MV1536/2021 9 AY.39.1.1 +NZL/21MV1298/2021 9 AY.39.1.1 +NZL/21MV1215/2021 9 AY.39.1.1 +England/NEWC-202DDC4/2021 27 AY.4 +BRA/ILMD2200125/2021 11 AY.101 +USA/MN-CDC-IBX711673689055/2021 2 AY.44 +BRA/BA-NVBS7150GENOV045616267287/2021 27 B.1.617.2 +BRA/PR-NVBS2685GENOV45371166115/2021 11 AY.101 +BRA/PR-NVBS4508GENOV827964198877/2021 11 AY.101 +BRA/RJ-NVBS6623GENOV828104289868/2021 1 AY.99.2 +BRA/ILMD2200096/2021 1 AY.99.2 +OY342278 27 AY.4 +OY382565 27 AY.4.2 +OY501555 27 AY.4 +NZL/21MV1387/2021 -1 AY.98 +NZL/21MV1933/2021 9 AY.39.1.1 +LAO/LOMWRU-1375/2021 3 AY.1 +IND/CWS_0011_49330959/2021 27 B.1.617.2 +BRA/RJ-NVBS5528GENOV827972782105/2021 1 AY.99.2 +LAO/LOMWRU-0826/2021 5 AY.85 +USA/SC-COVID21-5260/2021 22 AY.103 +DJI/NAMRU3_F397/2021 -1 AY.127 +England/ALDP-1633654/2021 27 AY.4 +England/MILK-179B06A/2021 27 AY.6 +England/PHEC-3J089J73/2021 27 AY.4 +IND/hospital_2_125/2021 27 B.1.617.2 +Scotland/QEUH-15D489C/2021 27 AY.4 +USA/CO-CDC-MMB09185280/2021 23 AY.25 +BRA/RJ-NVBS5406GENOV827934953444/2021 1 AY.99.2 +SouthAfrica/NHLS-UCT-GS-AE23/2021 26 AY.45 +NZL/21MV2839/2021 8 AY.39.1.1 +NZL/21MV2962/2021 8 AY.39.1.1 +NZL/21CH0644/2021 8 AY.39.1.1 +NZL/21CH0505/2021 9 AY.39.1.1 +NZL/21MV2031/2021 8 AY.39.1.1 +LAO/LOMWRU-1051/2021 3 AY.1 +DJI/NAMRU3_G907/2021 12 AY.100 +NZL/21MV1578/2021 9 AY.39.1.1 +OU517713 27 AY.4 +USA/MA-CDCBI-CRSP_O7KATHJL3WNZPUTM/2021 2 AY.44 +BGD/Laila_180/2021 27 B.1.617.2 +Scotland/SCOT-2944/2022 27 AY.4.2 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b/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts.tsv new file mode 100644 index 00000000..a65102bb --- /dev/null +++ b/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts.tsv @@ -0,0 +1,70 @@ +Nextclade_pango_collapsed t-sne_label_for_Nextclade_pango_collapsed count +AY.99.2 1 139 +B.1.617.2 27 241 +AY.4.2 27 29 +AY.29 16 41 +AY.4 27 241 +AY.3 18 44 +AY.3 15 13 +AY.101 11 32 +AY.45 26 26 +AY.43 21 98 +AY.33 27 1 +AY.39.1.1 9 82 +AY.44 2 80 +AY.98 -1 11 +AY.1 3 32 +AY.85 5 35 +AY.103 22 10 +AY.127 -1 11 +AY.6 27 22 +AY.25 23 40 +AY.39.1.1 8 58 +AY.100 12 21 +AY.122 14 27 +AY.46 -1 17 +AY.46 27 38 +AY.46.6 27 11 +B.1.617.2 -1 58 +AY.39 9 15 +AY.32 4 41 +AY.103 24 60 +AY.126 17 18 +AY.33 7 12 +AY.91 20 11 +AY.107 27 1 +AY.4 -1 19 +AY.34.1 10 14 +AY.39.1.1 -1 3 +AY.5 27 12 +AY.36 6 48 +AY.20 27 11 +AY.122 27 51 +B.1.617.2 10 5 +AY.23 13 18 +AY.25.1 23 18 +AY.98 27 7 +AY.103 -1 11 +AY.122 19 25 +AY.116 27 27 +AY.103 27 2 +AY.39 -1 14 +AY.120 25 15 +AY.122 -1 12 +AY.43 -1 4 +AY.46 10 1 +AY.119 27 11 +AY.99 0 11 +AY.107 -1 9 +AY.3 27 1 +B.1.617.2 12 2 +AY.25 -1 3 +AY.120 -1 3 +AY.91 -1 2 +AY.119 -1 2 +AY.4.2 -1 5 +AY.1 27 1 +AY.127 27 5 +B.1.617.2 7 1 +B.1.617.2 11 1 +AY.5 -1 2 diff --git a/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts_without_unclustered.tsv b/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts_without_unclustered.tsv new file mode 100644 index 00000000..7412e6b6 --- /dev/null +++ b/sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts_without_unclustered.tsv @@ -0,0 +1,53 @@ +Nextclade_pango_collapsed t-sne_label_for_Nextclade_pango_collapsed count +AY.99.2 1 139 +B.1.617.2 27 241 +AY.4.2 27 29 +AY.29 16 41 +AY.4 27 241 +AY.3 18 44 +AY.3 15 13 +AY.101 11 32 +AY.45 26 26 +AY.43 21 98 +AY.33 27 1 +AY.39.1.1 9 82 +AY.44 2 80 +AY.1 3 32 +AY.85 5 35 +AY.103 22 10 +AY.6 27 22 +AY.25 23 40 +AY.39.1.1 8 58 +AY.100 12 21 +AY.122 14 27 +AY.46 27 38 +AY.46.6 27 11 +AY.39 9 15 +AY.32 4 41 +AY.103 24 60 +AY.126 17 18 +AY.33 7 12 +AY.91 20 11 +AY.107 27 1 +AY.34.1 10 14 +AY.5 27 12 +AY.36 6 48 +AY.20 27 11 +AY.122 27 51 +B.1.617.2 10 5 +AY.23 13 18 +AY.25.1 23 18 +AY.98 27 7 +AY.122 19 25 +AY.116 27 27 +AY.103 27 2 +AY.120 25 15 +AY.46 10 1 +AY.119 27 11 +AY.99 0 11 +AY.3 27 1 +B.1.617.2 12 2 +AY.1 27 1 +AY.127 27 5 +B.1.617.2 7 1 +B.1.617.2 11 1 diff --git a/sars-cov-2-nextstrain/scripts/plot-single-clade-lineage-and-cluster-counts.py b/sars-cov-2-nextstrain/scripts/plot-single-clade-lineage-and-cluster-counts.py new file mode 100644 index 00000000..8ca45fee --- /dev/null +++ b/sars-cov-2-nextstrain/scripts/plot-single-clade-lineage-and-cluster-counts.py @@ -0,0 +1,28 @@ +import argparse +import altair as alt +import pandas as pd + + +if __name__ == "__main__": + parser = argparse.ArgumentParser() + parser.add_argument("--counts", required=True, help="path to TSV of counts by lineage and cluster") + parser.add_argument("--lineage-column", required=True, help="name of column in counts file for lineages") + parser.add_argument("--cluster-column", required=True, help="name of column in counts file for clusters") + parser.add_argument("--output", required=True, help="PNG dot plot of counts by lineage and cluster") + + args = parser.parse_args() + + df = pd.read_csv( + args.counts, + sep="\t", + ) + + chart = alt.Chart(df).mark_circle().encode( + x=alt.X(f"{args.lineage_column}:N", title="Pango lineage"), + y=alt.Y("{args.cluster_column}:N", title="Cluster from t-SNE"), + size="count:Q", + ).properties( + width=400, + height=400, + ) + chart.save(args.output, format="png", scale_factor=2.0) From b2c48d58058f25a6652a3283cb7a32db130c1d38 Mon Sep 17 00:00:00 2001 From: John Huddleston Date: Wed, 14 Aug 2024 13:46:23 -0700 Subject: [PATCH 3/8] Fix f-string bug that linting would have caught --- ...gle_clade_lineages_and_clusters_counts.png | Bin 79680 -> 90502 bytes 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figure, and adds references to this supplemental figure and the supplemental table of accuracy values which will eventually contain the accuracy values from this analysis. --- manuscript/cartography.tex | 15 +++++++-------- manuscript/cartography_supplement.tex | 7 +++++++ ...ingle-clade-lineages-and-clusters-counts.png} | Bin sars-cov-2-nextstrain/Snakefile | 4 ++-- 4 files changed, 16 insertions(+), 10 deletions(-) rename manuscript/figures/{sarscov2_single_clade_lineages_and_clusters_counts.png => sarscov2-single-clade-lineages-and-clusters-counts.png} (100%) diff --git a/manuscript/cartography.tex b/manuscript/cartography.tex index ef72abbe..c5bd65af 100644 --- a/manuscript/cartography.tex +++ b/manuscript/cartography.tex @@ -437,14 +437,13 @@ \subsection{SARS-CoV-2 clusters recapitulate broad genetic groups corresponding Unlike the Pango lineages in the early SARS-CoV-2 data, the lineages from the later data exhibited fewer pairwise genetic distances between samples in each lineage than samples in Nextstrain clades or any embedding cluster (Supplementary Fig.~S\ref{S_Fig_sarscov2_within_between_group_distances}). To understand whether t-SNE clusters could capture Pango-resolution genetic groups within a single Nextstrain clade, we evenly sampled approximately 2,000 sequences from a dominant Nextstrain clade with many Pango lineages, 21J (Delta), and identified clusters from a t-SNE embedding of those data. -Within the 1,992 sequences from 21J (Delta), we found 37 Pango lineages after collapsing lineages with fewer than 10 sequences into their parent lineages. -We found 44 t-SNE clusters representing 1,466 sequences with 526 sequences (26\%) assigned to the unclustered ``-1'' label. -The VI distance between all clusters (including the ``-1'' group) and the Pango lineages was 0.16. -This distance was consistent with the distance of 0.14 between t-SNE clusters and Pango lineages from both the full early and late SARS-CoV-2 datasets. -The VI distance between clusters without the -1 group and Pango lineages was 0.07, confirming that half of the distance between t-SNE clusters and Pango lineages above came from unclustered sequences. -Of the 35 Pango lineages with a t-SNE cluster, 21 lineages (60\%) had a single corresponding t-SNE cluster and 12 lineages (34\%) had 2 or 3 t-SNE clusters. -The remaining two lineages, AY.4 and B.1.617.2, had 5 and 10 t-SNE clusters, respectively. -Of the 44 t-SNE clusters, 29 clusters (66\%) had a single corresponding Pango lineage, 14 (32\%) mapped to two Pango lineages, and one cluster (43) mapped to 6 Pango lineages. +Within the 1,992 sequences from 21J (Delta), we found 38 Pango lineages after collapsing lineages with fewer than 10 sequences into their parent lineages. +We found 28 t-SNE clusters representing 1,806 sequences (91\%) with 186 sequences (9\%) assigned to the unclustered ``-1'' label (Supplementary Fig.~\ref{S_Fig_sarscov2_single_clade_embeddings_tsne_counts}). +The VI distance between Pango lineages and all clusters (including the unclustered group) was 0.17 (Supplementary Table~\ref{S_Table_optimal_cluster_parameters}). +This distance was consistent with the distance of 0.14 between Pango lineages and t-SNE clusters from both the full early and late SARS-CoV-2 datasets. +The VI distance between Pango lineages and clusters without the unclustered sequences was 0.13, confirming that one quarter of the distance between t-SNE clusters and Pango lineages above came from unclustered sequences. +Of the 38 Pango lineages with a t-SNE cluster, 30 lineages (79\%) had a single corresponding t-SNE cluster, seven lineages (18\%) had two or three t-SNE clusters, and one lineage (B.1.617.2) had five t-SNE clusters (Supplementary Fig.~\ref{S_Fig_sarscov2_single_clade_embeddings_tsne_counts}). +Of the 28 t-SNE clusters, 21 clusters (75\%) had a single corresponding Pango lineage, six (21\%) mapped to two or three Pango lineages, and one (cluster 27) mapped to 18 Pango lineages with most sequences from B.1.617.2 and AY.4. These results suggest that clusters from t-SNE embeddings can capture Pango-resolution genetic groups when analyzing sequences within a specific Nextstrain clade. \subsection{Distance-based embeddings reflect SARS-CoV-2 recombination events} diff --git a/manuscript/cartography_supplement.tex b/manuscript/cartography_supplement.tex index dfe6a21b..fe031839 100644 --- a/manuscript/cartography_supplement.tex +++ b/manuscript/cartography_supplement.tex @@ -209,6 +209,13 @@ \section*{Supplementary data} \caption{{\bf Number of late SARS-CoV-2 strains per combination of recombinant Pango lineage and t-SNE cluster where the count was at least 10 strains.}}\label{S_Fig_sarscov2_late_embeddings_tsne_recombinant_counts} \end{figure} +\begin{figure}[!h] +\includegraphics[width=\columnwidth]{figures/sarscov2-single-clade-lineages-and-clusters-counts.png} +\caption{{\bf Number of SARS-CoV-2 sequences from Nextstrain clade 21J (Delta) per combination of Pango lineage and t-SNE cluster showing the degree to which clusters of sequences from a single Nextstrain clade can capture Pango-resolution genetic groups.} + The size of each circle represents the number of sequences associated with each pair of t-SNE cluster and Pango lineage. + The cluster label ``-1'' represents sequences that could not be assigned to a cluster.}\label{S_Fig_sarscov2_single_clade_embeddings_tsne_counts} +\end{figure} + \begin{figure}[!h] \includegraphics[width=0.75\columnwidth]{figures/sarscov2-test-mds-reassortment-XE-BA1-BA2.png}\\ \includegraphics[width=0.75\columnwidth]{figures/sarscov2-test-mds-reassortment-XAS-BA2-BA5.png} diff --git a/manuscript/figures/sarscov2_single_clade_lineages_and_clusters_counts.png b/manuscript/figures/sarscov2-single-clade-lineages-and-clusters-counts.png similarity index 100% rename from manuscript/figures/sarscov2_single_clade_lineages_and_clusters_counts.png rename to manuscript/figures/sarscov2-single-clade-lineages-and-clusters-counts.png diff --git a/sars-cov-2-nextstrain/Snakefile b/sars-cov-2-nextstrain/Snakefile index 80e3dd6e..512e0101 100644 --- a/sars-cov-2-nextstrain/Snakefile +++ b/sars-cov-2-nextstrain/Snakefile @@ -21,7 +21,7 @@ rule sarscov2: "sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_without_unclustered_t-sne_for_Nextclade_pango_collapsed.csv", "sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts.tsv", "sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts_without_unclustered.tsv", - "manuscript/figures/sarscov2_single_clade_lineages_and_clusters_counts.png", + "manuscript/figures/sarscov2-single-clade-lineages-and-clusters-counts.png", rule sarscov2_download_filtered_metadata: output: @@ -1192,7 +1192,7 @@ rule sarscov2_plot_lineages_and_clusters_for_single_clade: input: counts="sars-cov-2-nextstrain/results/single_clade/lineages_and_clusters_counts.tsv", output: - counts_plot="manuscript/figures/sarscov2_single_clade_lineages_and_clusters_counts.png", + counts_plot="manuscript/figures/sarscov2-single-clade-lineages-and-clusters-counts.png", conda: "../cartography.yml" params: lineage_column="Nextclade_pango_collapsed", From 0b4d099fce1d4238382341927fee7144982afc7b Mon Sep 17 00:00:00 2001 From: John Huddleston Date: Wed, 14 Aug 2024 14:11:58 -0700 Subject: [PATCH 5/8] Include within-clade cluster accuracy in table Adds the within-clade cluster accuracy file to the overall accuracy table. The next commit(s) will update the table itself. --- manuscript/Snakefile | 3 +++ 1 file changed, 3 insertions(+) diff --git a/manuscript/Snakefile b/manuscript/Snakefile index 1713a71b..bbf1cfc3 100644 --- a/manuscript/Snakefile +++ b/manuscript/Snakefile @@ -69,6 +69,7 @@ rule concat_HDBSCAN_tables: "sars-cov-2-nextstrain/results/optimal_cluster_accuracy_and_parameters_for_Nextclade_pango_collapsed.csv", "sars-cov-2-nextstrain-2022-2023/results/full_HDBSCAN_metadata_for_Nextstrain_clade.csv", "sars-cov-2-nextstrain-2022-2023/results/full_HDBSCAN_metadata_for_Nextclade_pango_collapsed.csv", + "sars-cov-2-nextstrain/results/single_clade/cluster_accuracy_t-sne_for_Nextclade_pango_collapsed.csv", ] output: fullKDE = "manuscript/tables/total_HDBSCAN_table.csv", @@ -83,6 +84,7 @@ rule concat_HDBSCAN_tables: "early SARS-CoV-2 (2020-2022)", "late SARS-CoV-2 (2022-2023)", "late SARS-CoV-2 (2022-2023)", + "SARS-CoV-2 21J (Delta) only", ], genetic_group_types = [ "Nextstrain clade", @@ -93,6 +95,7 @@ rule concat_HDBSCAN_tables: "Pango", "Nextstrain clade", "Pango", + "Pango", ], conda: "../cartography.yml" shell: From a3274781ab28674a90c0f3137f6dd0a74cb09753 Mon Sep 17 00:00:00 2001 From: John Huddleston Date: Wed, 14 Aug 2024 14:21:29 -0700 Subject: [PATCH 6/8] Add within-clade accuracy to supplemental table Adds the within-clade accuracy for 21J (Delta) SARS-CoV-2 sequences to supplemental table 1 after fixing a bug in the script that creates that table that assumes all inputs will have the same number of rows. --- manuscript/tables/accuracy_table.tex | 1 + manuscript/tables/total_HDBSCAN_table.csv | 1 + notebooks/scripts/concat_KDE_tables.py | 5 +++-- 3 files changed, 5 insertions(+), 2 deletions(-) diff --git a/manuscript/tables/accuracy_table.tex b/manuscript/tables/accuracy_table.tex index fab7fdf9..4e371703 100644 --- a/manuscript/tables/accuracy_table.tex +++ b/manuscript/tables/accuracy_table.tex @@ -42,5 +42,6 @@ & & MDS & 18 & 0.37 & \\ & & genetic & 15 & 0.43 & \\ & & PCA & 7 & 0.46 & \\ + SARS-CoV-2 21J (Delta) only & Pango & t-SNE & 28 & 0.17 & \\ \botrule \end{tabular} diff --git a/manuscript/tables/total_HDBSCAN_table.csv b/manuscript/tables/total_HDBSCAN_table.csv index 60f1f469..9b3867ed 100644 --- a/manuscript/tables/total_HDBSCAN_table.csv +++ b/manuscript/tables/total_HDBSCAN_table.csv @@ -39,3 +39,4 @@ 37,MDS,0.37,18,,, 38,genetic,0.43,15,,, 39,PCA,0.46,7,,, +40,t-SNE,0.17,28,,SARS-CoV-2 21J (Delta) only,Pango diff --git a/notebooks/scripts/concat_KDE_tables.py b/notebooks/scripts/concat_KDE_tables.py index db6cebbe..d35fcf55 100644 --- a/notebooks/scripts/concat_KDE_tables.py +++ b/notebooks/scripts/concat_KDE_tables.py @@ -42,8 +42,9 @@ df = df.loc[:, [column for column in columns_to_use if column != "distance_threshold"]].copy() df["distance_threshold"] = "" - df["Pathogen Dataset"] = [pathogen] + [""] * (len(embedding_name_by_abbreviation) - 1) - df["Genetic Group Type"] = [genetic_group_type] + [""] * (len(embedding_name_by_abbreviation) - 1) + number_of_methods = df["method"].drop_duplicates().shape[0] + df["Pathogen Dataset"] = [pathogen] + [""] * (number_of_methods - 1) + df["Genetic Group Type"] = [genetic_group_type] + [""] * (number_of_methods - 1) tables.append(df) df = pd.concat(tables, ignore_index=True) From adbab82ffd93fa705491f53a09433461f2750b1f Mon Sep 17 00:00:00 2001 From: John Huddleston Date: Wed, 14 Aug 2024 14:25:23 -0700 Subject: [PATCH 7/8] Small text edit --- manuscript/cartography.tex | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/manuscript/cartography.tex b/manuscript/cartography.tex index c5bd65af..9b366b13 100644 --- a/manuscript/cartography.tex +++ b/manuscript/cartography.tex @@ -437,7 +437,7 @@ \subsection{SARS-CoV-2 clusters recapitulate broad genetic groups corresponding Unlike the Pango lineages in the early SARS-CoV-2 data, the lineages from the later data exhibited fewer pairwise genetic distances between samples in each lineage than samples in Nextstrain clades or any embedding cluster (Supplementary Fig.~S\ref{S_Fig_sarscov2_within_between_group_distances}). To understand whether t-SNE clusters could capture Pango-resolution genetic groups within a single Nextstrain clade, we evenly sampled approximately 2,000 sequences from a dominant Nextstrain clade with many Pango lineages, 21J (Delta), and identified clusters from a t-SNE embedding of those data. -Within the 1,992 sequences from 21J (Delta), we found 38 Pango lineages after collapsing lineages with fewer than 10 sequences into their parent lineages. +Within the 1,992 sequences sampled from 21J (Delta), we found 38 Pango lineages after collapsing lineages with fewer than 10 sequences into their parent lineages. We found 28 t-SNE clusters representing 1,806 sequences (91\%) with 186 sequences (9\%) assigned to the unclustered ``-1'' label (Supplementary Fig.~\ref{S_Fig_sarscov2_single_clade_embeddings_tsne_counts}). The VI distance between Pango lineages and all clusters (including the unclustered group) was 0.17 (Supplementary Table~\ref{S_Table_optimal_cluster_parameters}). This distance was consistent with the distance of 0.14 between Pango lineages and t-SNE clusters from both the full early and late SARS-CoV-2 datasets. From e4b95614ea7715fbf8054f08b6c5d14be12e0c69 Mon Sep 17 00:00:00 2001 From: John Huddleston Date: Wed, 14 Aug 2024 14:35:25 -0700 Subject: [PATCH 8/8] Clarify the discussion re: within-clade results --- manuscript/cartography.tex | 5 +++-- 1 file changed, 3 insertions(+), 2 deletions(-) diff --git a/manuscript/cartography.tex b/manuscript/cartography.tex index 9b366b13..85a276b6 100644 --- a/manuscript/cartography.tex +++ b/manuscript/cartography.tex @@ -444,7 +444,7 @@ \subsection{SARS-CoV-2 clusters recapitulate broad genetic groups corresponding The VI distance between Pango lineages and clusters without the unclustered sequences was 0.13, confirming that one quarter of the distance between t-SNE clusters and Pango lineages above came from unclustered sequences. Of the 38 Pango lineages with a t-SNE cluster, 30 lineages (79\%) had a single corresponding t-SNE cluster, seven lineages (18\%) had two or three t-SNE clusters, and one lineage (B.1.617.2) had five t-SNE clusters (Supplementary Fig.~\ref{S_Fig_sarscov2_single_clade_embeddings_tsne_counts}). Of the 28 t-SNE clusters, 21 clusters (75\%) had a single corresponding Pango lineage, six (21\%) mapped to two or three Pango lineages, and one (cluster 27) mapped to 18 Pango lineages with most sequences from B.1.617.2 and AY.4. -These results suggest that clusters from t-SNE embeddings can capture Pango-resolution genetic groups when analyzing sequences within a specific Nextstrain clade. +These results suggest that clusters from t-SNE embeddings can capture more Pango-resolution genetic groups by analyzing sequences within a specific Nextstrain clade. \subsection{Distance-based embeddings reflect SARS-CoV-2 recombination events} @@ -492,7 +492,8 @@ \subsection{Limitations of methods and analysis} Despite the promise of these simple methods to answer important public health questions about human pathogenic viruses, these methods and our analyses suffer from inherent limitations. The lack of an underlying biological model is both a strength and the clearest limitation of the dimensionality reduction methods we considered here. -For example, embeddings of SARS-CoV-2 genomes cannot capture the same fine-grained genetic resolution as Pango lineage annotations. +For example, embeddings of SARS-CoV-2 genomes that represent the complete circulating diversity at a given time cannot capture the same fine-grained genetic resolution as Pango lineage annotations. +Only t-SNE clusters of SARS-CoV-2 genomes within a single Nextstrain clade get close to defining Pango-resolution genetic groups. Each method provides only a few parameters to tune its embeddings and these parameters have little effect on the qualitative outcome. In maintaining a linear relationship between Euclidean and genetic distances, MDS sacrifices the ability to form more accurate genetic clusters for viruses with large genomes like SARS-CoV-2. Neither t-SNE nor UMAP maintain a linear relationship between pairwise Euclidean and genetic distances across the observed range of genetic distances.