From 3bdaf51034cce2fa0b07a5905edefa290d8660a0 Mon Sep 17 00:00:00 2001 From: Tal Scully Date: Wed, 7 May 2025 17:02:11 -0400 Subject: [PATCH 01/14] Added code for generating inferred hematopoietic hierarchy coarse-grain tree --- .gitignore | 1 + .../README.md | 6 +- .../differentiation_graph.py | 300 ++++++++++++++++++ .../differentiation_graph_env.txt | 59 ++++ .../cell_type_graph_1.pdf | Bin 0 -> 15317 bytes .../cell_type_graph_2.pdf | Bin 0 -> 4915 bytes .../cell_type_graph_3.pdf | Bin 0 -> 8335 bytes .../cell_type_graph_adjacency_matrix.pdf | Bin 0 -> 27111 bytes .../connections_heatmap_all_to_all.pdf | Bin 0 -> 37613 bytes .../connections_heatmap_prog_to_mature.pdf | Bin 0 -> 23004 bytes .../edge_weight_distribution.pdf | Bin 0 -> 16085 bytes .../mki67_expr_per_cell_type.png | Bin 0 -> 622075 bytes .../nodes_without_edges.txt | 8 + 13 files changed, 373 insertions(+), 1 deletion(-) create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph.py create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_env.txt create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/cell_type_graph_1.pdf create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/cell_type_graph_2.pdf create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/cell_type_graph_3.pdf create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/cell_type_graph_adjacency_matrix.pdf create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/connections_heatmap_all_to_all.pdf create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/connections_heatmap_prog_to_mature.pdf create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/edge_weight_distribution.pdf create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/mki67_expr_per_cell_type.png create mode 100644 Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_output/nodes_without_edges.txt diff --git a/.gitignore b/.gitignore index dac829b..c4e167a 100644 --- a/.gitignore +++ b/.gitignore @@ -5,3 +5,4 @@ __pycache__/ Scully_Ciona_blood_2025/data/* !Scully_Ciona_blood_2025/data/README.md Scully_Ciona_blood_2025/scRNA-seq_analysis/go_enrichment_analysis/go_enrichment_analysis_output +Scully_Ciona_blood_2025/scRNA-seq_analysis/go_enrichment_analysis/differentiation_graph_output diff --git a/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/README.md b/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/README.md index 18bf624..152ae22 100644 --- a/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/README.md +++ b/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/README.md @@ -1 +1,5 @@ -Scripts coming soon... \ No newline at end of file +The scripts in this folder were used to generate a hypothesized hematopoietic hierarchy for _C. robsuta_, shown in **Fig. 4c** of the paper. See methods section "Differentiation Hierarchy Analysis". + +The files are: +- `differentiation_graph.py` is the python script that uses the scRNA-seq data to generate a hypothesized hierarchy. The script outputs a folder `differentiation_graph_output/` containing plots of the inferred hierarchy (graph, adjacency matrix), a text file listing cell states not connected to other nodes in the graph, as well as miscellaneous supporting plots. +- `differentiation_graph_env.txt` contains information on the conda environment used, saved to a text file (by running `conda list --explicit > differentiation_graph_env.txt`). diff --git a/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph.py b/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph.py new file mode 100644 index 0000000..93e952b --- /dev/null +++ b/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph.py @@ -0,0 +1,300 @@ +import numpy as np +import pandas as pd +import matplotlib.pyplot as plt +import seaborn as sns +import scanpy as sc +import networkx as nx +import sys +import os +from tqdm import tqdm + +# Change this path to point to folder containing gene_hf.py +# This imports dictionaries and functions for easily converting gene ids +path_to_repo_dir = os.path.dirname(os.path.dirname(os.path.dirname(os.path.abspath(__file__)))) +sys.path.append(os.path.join(path_to_repo_dir, 'helper_functions')) +import scrna_helper_functions as hf + +plt.style.use('tal_paper_spine') + +# Set random seed +np.random.seed(seed = 0) + +# ============================================================================ +# IMPORT DATA + +# Import counts matrix +adata = sc.read_h5ad(hf.path.Crob_adata_file) + +# Set up output folder +out_path = os.path.join(os.path.dirname(os.path.abspath(__file__)), + 'differentiation_graph_output') +if not os.path.exists(out_path): os.mkdir(out_path) + +# ============================================================================ +# DEFINE CYCLING PROGENITOR CELL STATES +# Cell states are defined to be cycling if their median expression of MKI67 +# is above 0. + +percentile = 50 # 50th percentile = median +percentile_dict = {} +for cell_type in adata.obs['cell_type'].cat.categories: + adata_subset = adata[adata.obs['cell_type'] == cell_type] + percentile_dict[cell_type] = np.percentile( + np.array(adata_subset.raw[:, hf.human2ciona['MKI67']].X.todense()), + percentile) + +with plt.style.context('tal_light'): + f = plt.figure(figsize=(10, 8)) + ax = plt.subplot(2, 2, 1) + sc.pl.umap(adata, color=hf.human2ciona['MKI67'], show=False, ax=ax) + ax = plt.subplot(2, 2, 2) + sc.pl.umap(adata, color='cell_type', show=False, ax=ax, + legend_loc='on data') + + ax = plt.subplot(2, 1, 2) + plt.bar([c for c in percentile_dict], + [percentile_dict[c] for c in percentile_dict]) + plt.xticks(rotation=90) + plt.ylabel(f'{percentile}th percentile expression') + + plt.tight_layout() + plt.savefig(os.path.join(out_path, 'mki67_expr_per_cell_type.png'), + dpi=200) + plt.close() + +# ============================================================================ +# BUILD A COARSE-GRAIN WEIGHTED GRAPH FROM THE SINGLE-CELL GRAPH + +# From Wagner et al. 2018 methods section for building coarse-grain graph: +""" +Construction of Coarse-Grained Graphs +A coarse-graining procedure to abstract the major features of the single-cell +graph was performed as follows. First, single-cell nodes belonging to the same +annotated tSNE cluster ID were collapsed into a single state node. Edges +between each pair of state nodes were then weighted by calculating the norm +index of original shared single-cell edges (i.e. the ratio of shared single- +cell edges to the total number of outgoing edges for that node pair). State +edges were then discarded if they received a norm index weight < 0.01. +Finally, a spanning tree was traced through the weighted edges as follows. +Beginning with the final timepoint, edges for all nodes were ranked according +to weight. Edges then were then removed recursively, starting with the weakest +edges, unless doing so would increase the total number of graph connected +components. This process was then repeated for each timepoint. The resulting +spanning tree connects all nodes to a single 4hpf “root” node defined by all +cells of the first timepoint. +""" + +# ------------------------------------ +# Get cell-cell edges that we care about + +# Convert the single-cell k-nearest neighbors graph sparse matrix to a +# NetworkX graph +G = nx.from_scipy_sparse_matrix(adata.obsp['connectivities']) + +# Filter cell edges to retain only those connected to EXACTLY 1 progenitor +# cell_edges = [] +# for i in selected_cells: +# neighbors = list(G.neighbors(i)) +# for j in neighbors: +# # if j not in selected_cells: +# cell_edges.append((i, j)) + +cell_edges = list(G.edges) + +# ------------------------------------ +# Count cell edges to create a weighted cell type graph + +cell_type_list = adata.obs['cell_type'].cat.categories +cLRP_list = [c for c in cell_type_list if 'cLRP' in c] +prog_list = [c for c in cell_type_list if c[0] == 'c'] +other_list = [c for c in cell_type_list if c not in prog_list] +prog_to_mature = pd.DataFrame(0, index=other_list, columns=prog_list) +all_to_all = pd.DataFrame(0, index=cell_type_list, columns=cell_type_list) + +for (i, j) in tqdm(cell_edges): + cell1 = adata.obs.iloc[i]['cell_type'] + cell2 = adata.obs.iloc[j]['cell_type'] + if cell1 != cell2: + # Edges between progenitor and non-progenitor cell types + if (cell1 in prog_list) and (cell2 not in prog_list): + prog_to_mature.loc[cell2, cell1] += 1 + elif (cell2 in prog_list) and (cell1 not in prog_list): + prog_to_mature.loc[cell1, cell2] += 1 + # Edges between cLRP and other cell types + else: + all_to_all.loc[cell1, cell2] += 1 + all_to_all.loc[cell2, cell1] += 1 + +# Calculate normalized edge counts +prog_to_mature_norm = prog_to_mature.copy() +total_edges_row = prog_to_mature.sum(axis=1) +total_edges_col = prog_to_mature.sum(axis=0) +for row in prog_to_mature_norm.index: + for col in prog_to_mature_norm.columns: + prog_to_mature_norm.loc[row, col] = ( + prog_to_mature.loc[row, col] + / np.sqrt(total_edges_row[row] * total_edges_col[col]) + ) +all_to_all_norm = all_to_all.copy() +total_edges_row = all_to_all.sum(axis=1) +total_edges_col = all_to_all.sum(axis=0) +for row in all_to_all_norm.index: + for col in all_to_all_norm.columns: + all_to_all_norm.loc[row, col] = ( + all_to_all.loc[row, col] + / np.sqrt(total_edges_row[row] * total_edges_col[col]) + ) + +# ------------------------------------ +# Plot as heatmap + +graph_dict = { + 'prog_to_mature': prog_to_mature_norm, + 'all_to_all': all_to_all_norm, +} + +for graph_name in graph_dict: + norm_index = graph_dict[graph_name] + + # Reorder for plotting + marginals_col = norm_index.max(axis=0) + prog_order = norm_index.columns[np.argsort(marginals_col)[::-1]] + marginals_row = norm_index.max(axis=1) + other_order = norm_index.index[np.argsort(marginals_row)[::-1]] + + f = plt.figure(figsize=(1.126 + 0.1*norm_index.shape[1], + 0.6297 + 0.1*norm_index.shape[0])) + sns.heatmap(norm_index.loc[other_order, prog_order], + cmap='rocket_r', xticklabels=True, yticklabels=True, + cbar_kws={'label': 'Normalized edge counts'}) + plt.tight_layout() + plt.savefig(os.path.join(out_path, f'connections_heatmap_{graph_name}.pdf')) + plt.close() + +# ============================================================================ +# REMOVING LOW-WEIGHT EDGES AND GET MAXIMUM SPANNING TREE + +# Graph of cell types +G = nx.Graph() +G.add_nodes_from(cell_type_list) +G.nodes['cMPP']['group'] = 'cMPP' +for n in cLRP_list: + G.nodes[n]['group'] = 'cLRP' +for n in other_list: + G.nodes[n]['group'] = 'mature' + +# Set edge weight threshold +f = plt.figure(figsize=(3, 4)) +percentage = 30 +for i, graph_name in enumerate(graph_dict): + norm_index = graph_dict[graph_name] + threshold = (percentage/100) * norm_index.max().max() + + freq, bins = np.histogram(norm_index.values.flatten(), bins=10) + ax = plt.subplot(2, 1, i+1) + plt.bar(bins[:-1], freq, width=np.diff(bins), color='#888888') + plt.axvline(x=threshold,label=f'Threshold = {percentage}%\nof max weight') + plt.yscale('log') + plt.ylabel('Frequency') + plt.legend() + plt.title(graph_name) +# Finish plot +plt.xlabel('Edge weight') +plt.tight_layout() +plt.savefig(os.path.join(out_path, 'edge_weight_distribution.pdf')) +plt.close() + +# Add edges to grpah object +graph_edges = [] +for node1 in G.nodes: + for node2 in G.nodes: + if node1 != node2: + # Weights from each graph, set to zero if below threshold + weight1 = all_to_all_norm.loc[node1, node2] + if weight1 < ((percentage/100) * all_to_all_norm.max().max()): + weight1 = 0 + if (node2 in prog_list) and (node1 not in prog_list): + weight2 = prog_to_mature_norm.loc[node1, node2] + if weight2 < ((percentage/100) * prog_to_mature_norm.max().max()): + weight2 = 0 + elif (node1 in prog_list) and (node2 not in prog_list): + weight2 = prog_to_mature_norm.loc[node2, node1] + if weight2 < ((percentage/100) * prog_to_mature_norm.max().max()): + weight2 = 0 + else: + weight2 = 0 + + # Take max of weights from each graph + weight = max(weight1, weight2) + if weight != 0: + if (node2, node1, {'weight': weight}) not in graph_edges: + graph_edges.append((node1, node2, {'weight': weight})) +G.add_edges_from(graph_edges) + +# Save nodes without edges, then remove them +with open(os.path.join(out_path, 'nodes_without_edges.txt'), 'w') as f: + f.write('\n'.join(list(nx.isolates(G)))) +G.remove_nodes_from(list(nx.isolates(G))) + +# Take the maximum spanning tree +G = nx.maximum_spanning_tree(G, weight='weight') + +# ============================================================================ +# VISUALIZE COARSE-GRAIN TREE + +fig_len = { + 0: 2.25, + 1: 0.5, + 2: 0.7, +} +for i, node_set in enumerate(list(nx.connected_components(G))): + subG = G.subgraph(node_set).copy() + + # Plot as a tree + # (Have to rename nodes to not get an error) + mapping = {node: f"node_{i}" for i, node in enumerate(subG.nodes)} + mapping_reverse = {mapping[node]: node for node in mapping} + subG = nx.relabel_nodes(subG, mapping) + pos = nx.drawing.nx_pydot.graphviz_layout(subG, prog="neato") + subG = nx.relabel_nodes(subG, mapping_reverse) + pos_renamed = {mapping_reverse[node]: pos[node] for node in mapping_reverse} + + # Step 3: Draw the graph + edge_weights = nx.get_edge_attributes(subG, 'weight') + color_map = {'cMPP': '#d94a34', 'cLRP': '#c5a801', 'mature': '#7db9d1'} + node_colors = [color_map[subG.nodes[node]['group']] for node in subG.nodes()] + f = plt.figure(figsize=(fig_len[i], fig_len[i])) + nx.draw( + subG, + pos_renamed, + with_labels=True, + node_size=350, + node_color=node_colors, + edge_color="gray", + font_size=7, + width=[7*weight for weight in edge_weights.values()], + labels={c: c.replace(' (', '\n(') for c in subG.nodes}, + alpha=1, + ) + f.savefig(os.path.join(out_path, f'cell_type_graph_{i+1}.pdf')) + plt.close() + +# ------------------------------------ +# Plot as heatmap + +df = pd.DataFrame(nx.adjacency_matrix(G).todense(), index=G.nodes(), + columns=G.nodes()) + +# Reorder for plotting +marginals_col = df.max(axis=0) +prog_order = df.columns[np.argsort(marginals_col)[::-1]] +marginals_row = df.max(axis=1) +other_order = df.index[np.argsort(marginals_row)[::-1]] + +f = plt.figure(figsize=(1.126 + 0.1*df.shape[1], 0.6297 + 0.1*df.shape[0])) +sns.heatmap(df.loc[other_order, prog_order], + cmap='rocket_r', xticklabels=True, yticklabels=True, + cbar_kws={'label': 'norm index'}) +plt.tight_layout() +plt.savefig(os.path.join(out_path, f'cell_type_graph_adjacency_matrix.pdf')) +plt.close() diff --git a/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_env.txt b/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_env.txt new file mode 100644 index 0000000..98583e3 --- /dev/null +++ b/Scully_Ciona_blood_2025/scRNA-seq_analysis/differentiation_hierarchy_analysis/differentiation_graph_env.txt @@ -0,0 +1,59 @@ +# This file may be used to create an environment using: +# $ conda create --name --file +# platform: osx-64 +@EXPLICIT +https://conda.anaconda.org/conda-forge/osx-64/ca-certificates-2024.8.30-h8857fd0_0.conda +https://conda.anaconda.org/conda-forge/osx-64/jpeg-9e-hb7f2c08_3.conda +https://repo.anaconda.com/pkgs/main/osx-64/libcxx-10.0.0-1.conda +https://repo.anaconda.com/pkgs/main/osx-64/libsodium-1.0.18-h1de35cc_0.conda +https://conda.anaconda.org/conda-forge/osx-64/libwebp-base-1.1.0-hbcb3906_3.tar.bz2 +https://repo.anaconda.com/pkgs/main/osx-64/xz-5.2.5-h1de35cc_0.conda +https://repo.anaconda.com/pkgs/main/osx-64/zlib-1.2.11-h1de35cc_3.conda +https://repo.anaconda.com/pkgs/main/osx-64/libffi-3.3-hb1e8313_2.conda +https://conda.anaconda.org/conda-forge/osx-64/libpng-1.6.37-h7cec526_2.tar.bz2 +https://conda.anaconda.org/conda-forge/osx-64/lz4-c-1.9.2-h4a8c4bd_1.tar.bz2 +https://repo.anaconda.com/pkgs/main/osx-64/ncurses-6.2-h0a44026_1.conda +https://conda.anaconda.org/conda-forge/osx-64/openssl-1.1.1w-h8a1eda9_0.conda 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