diff --git a/.gitignore b/.gitignore index c3b28032..79719dee 100644 --- a/.gitignore +++ b/.gitignore @@ -70,6 +70,8 @@ instance/ # Sphinx documentation docs/build/ +docs/source/auto_examples/ +output_files/ # PyBuilder target/ @@ -128,6 +130,9 @@ venv.bak/ .dmypy.json dmypy.json +# ruff +.ruff_cache/ + # Pyre type checker .pyre/ diff --git a/docs/source/.DS_Store b/docs/source/.DS_Store deleted file mode 100644 index 4e51ff10..00000000 Binary files a/docs/source/.DS_Store and /dev/null differ diff --git a/docs/source/index.rst b/docs/source/index.rst index 4de749ec..f228556d 100644 --- a/docs/source/index.rst +++ b/docs/source/index.rst @@ -17,7 +17,14 @@ from scratch, you may experience some differences in terms of retrieved values o Currently, available features broadly correspond to the default settings of DESeq2 (v1.34.0) for single-factor and multi-factor analysis (with categorical or continuous factors) using Wald tests, with an optional `apeGLM `_ LFC shrinkage step -:cite:p:`zhu2019heavy`. We plan to implement more in the near future. In case there is a feature you would particularly +:cite:p:`zhu2019heavy`. + +PyDESeq2 also supports `pytximport `_-derived +normalization factors :cite:p:`kuehlGeneCountEstimation2024b`, enabling accurate differential expression analysis +with transcript-level quantification data from tools like Salmon, Kallisto, or RSEM. When explicitly enabled, +this feature accounts for gene length differences between samples due to differential isoform usage. + +We plan to implement more features in the near future. In case there is a feature you would particularly like to be implemented, feel free to open an issue on GitHub. diff --git a/docs/source/refs.bib b/docs/source/refs.bib index a579d2d8..25f158fc 100644 --- a/docs/source/refs.bib +++ b/docs/source/refs.bib @@ -30,4 +30,16 @@ @article{zhu2019heavy year={2019}, doi={10.1093/bioinformatics/bty895}, publisher={Oxford University Press} -} \ No newline at end of file +} + +@article{kuehlGeneCountEstimation2024b, + title = {Gene Count Estimation with Pytximport Enables Reproducible Analysis of Bulk {{RNA}} Sequencing Data in {{Python}}}, + author = {Kuehl, Malte and Wong, Milagros N and Wanner, Nicola and Bonn, Stefan and Puelles, Victor G}, + year = {2024}, + journal = {Bioinformatics}, + volume = {40}, + number = {12}, + pages = {btae700}, + doi = {10.1093/bioinformatics/btae700}, + publisher = {Oxford University Press} +} diff --git a/examples/plot_pytximport_example.py b/examples/plot_pytximport_example.py new file mode 100644 index 00000000..a7735d74 --- /dev/null +++ b/examples/plot_pytximport_example.py @@ -0,0 +1,164 @@ +""" +pytximport Integration Example +============================== + +This example demonstrates how to use PyDESeq2 with pytximport-derived data +for differential expression analysis with transcript-level quantification data. + +pytximport enables the import of transcript-level abundance estimates from +popular RNA-seq quantification tools like Salmon, Kallisto, or RSEM into +PyDESeq2, while accounting for gene length differences between samples +due to differential isoform usage. +""" + +import anndata as ad +import matplotlib.pyplot as plt +import numpy as np + +from pydeseq2.dds import DeseqDataSet +from pydeseq2.ds import DeseqStats + +# %% +# Read AnnData object +# ------------------- +# +# Make sure that you have rounded the counts in .X to integers, as DESeq2 requires +# integer counts and that pytximport was used with counts_from_abundance=None +# (raw counts) to generate the AnnData object. + +adata = ad.read_h5ad("../tests/data/pytximport/test_pytximport.h5ad") + +# %% +# Standard usage without pytximport +# --------------------------------- +# +# By default, PyDESeq2 uses standard size factor normalization. + +dds_standard = DeseqDataSet(adata=adata, design="~condition") +dds_standard.fit_size_factors() +print(f"Standard normalization: {'normalization_factors' not in dds_standard.obsm}") + +# %% +# Explicit pytximport mode +# ------------------------ +# +# To use pytximport normalization factors, explicitly set from_pytximport=True. + +dds_explicit = DeseqDataSet(adata=adata, design="~condition", from_pytximport=True) +dds_explicit.fit_size_factors() +print(f"Pytximport normalization: {'normalization_factors' in dds_explicit.obsm}") + +# %% +# Size factor estimation with normalization factors +# ------------------------------------------------- +# +# When pytximport data is used, PyDESeq2 computes normalization factors +# that account for both library size and gene length differences. + +dds_explicit.fit_size_factors() + +print(f"Size factors computed: {'size_factors' in dds_explicit.obs}") +print(f"Normalization factors computed: {'normalization_factors' in dds_explicit.obsm}") + +# Check normalization factors properties +norm_factors = dds_explicit.obsm["normalization_factors"] +size_factors = dds_explicit.obs["size_factors"] + +print(f"Normalization factors shape: {norm_factors.shape}") +print(f"Size factors shape: {size_factors.shape}") +print(f"Size factors: {size_factors.values}") +print(f"Norm factors range: {norm_factors.min():.3f} - {norm_factors.max():.3f}") + +# %% +# Plot distribution of normalization factors +# ------------------------------------------ +# +# Normalization factors should vary around 1, reflecting gene length +# differences between samples. +plt.figure(figsize=(8, 5)) +plt.hist( + norm_factors.flatten(), + bins=list(np.arange(0.5, 1.6, 0.1).astype(float)), + color="skyblue", + edgecolor="black", + alpha=0.7, +) +plt.axvline(x=1, color="red", linestyle="--", linewidth=2, label="x = 1") +plt.title("Distribution of Normalization Factors") +plt.xlabel("Normalization Factor") +plt.ylabel("Frequency") +plt.legend() +plt.tight_layout() +plt.show() + +# %% +# Full differential expression analysis +# ------------------------------------- +# +# The complete DESeq2 pipeline works seamlessly with pytximport data. + +# Run the full DESeq2 pipeline +dds_explicit.deseq2() + +# Perform statistical testing +stat_res = DeseqStats(dds_explicit, contrast=["condition", "ko", "wt"]) +stat_res.summary() +results_df = stat_res.results_df + +print("Differential expression results:") +print(f"Total genes analyzed: {len(results_df)}") +print(f"Significant genes (padj < 0.05): {sum(results_df['padj'] < 0.05)}") + +# Display top results +print("\nTop 5 most significant genes:") +top_results = results_df.sort_values("padj").head() +print(top_results[["baseMean", "log2FoldChange", "pvalue", "padj"]]) +stat_res.plot_MA() + +# %% +# Differential expression analysis without normalization factors +# -------------------------------------------------------------- +# +# Compare results with standard DESeq2 normalization (without length correction). + +# Read standard AnnData (generated with counts_from_abundance="length_scaled_tpm") +adata_standard = ad.read_h5ad( + "../tests/data/pytximport/test_pytximport_length_scaled.h5ad" +) + +dds_standard = DeseqDataSet(adata=adata_standard, design="~condition") +dds_standard.deseq2() + +stat_res_standard = DeseqStats(dds_standard, contrast=["condition", "ko", "wt"]) +stat_res_standard.summary() +results_standard = stat_res_standard.results_df +print("\nTop 5 most significant genes:") +top_results = results_standard.sort_values("padj").head() +print(top_results[["baseMean", "log2FoldChange", "pvalue", "padj"]]) +stat_res_standard.plot_MA() + +# %% +# Comparison with standard normalization +# -------------------------------------- +# +# Compare results with standard DESeq2 normalization (without length correction). +pytximport_sig = sum(results_df["padj"] < 0.05) +standard_sig = sum(results_standard["padj"] < 0.05) + +print("\nComparison of significant genes (padj < 0.05):") +print(f"pytximport normalization: {pytximport_sig}") +print(f"Standard normalization: {standard_sig}") + +# %% +# Variance Stabilizing Transformation (VST) +# ----------------------------------------- +# +# VST also works with pytximport normalization factors. + +dds_vst = DeseqDataSet(adata=adata, design="~condition", from_pytximport=True) +dds_vst.vst() + +vst_counts = dds_vst.layers["vst_counts"] +print("\nVST transformation completed") +print(f"VST counts shape: {vst_counts.shape}") +print(f"VST counts range: {vst_counts.min():.3f} - {vst_counts.max():.3f}") diff --git a/pydeseq2/dds.py b/pydeseq2/dds.py index cfff35a3..6a8e90c7 100644 --- a/pydeseq2/dds.py +++ b/pydeseq2/dds.py @@ -16,6 +16,7 @@ from pydeseq2.inference import Inference from pydeseq2.preprocessing import deseq2_norm_fit from pydeseq2.preprocessing import deseq2_norm_transform +from pydeseq2.preprocessing import estimate_norm_factors from pydeseq2.utils import dispersion_trend from pydeseq2.utils import make_scatter from pydeseq2.utils import mean_absolute_deviation @@ -135,6 +136,14 @@ class DeseqDataSet(ad.AnnData): longer necessary after they are used during deseq2 run, such as Cook's distances. (default: False) + from_pytximport : bool + Whether the data comes from pytximport and should use length-corrected + normalization factors instead of simple size factors. Must be explicitly + set to True to enable pytximport support. When enabled, will validate the + presence of "length" in obsm and "counts_from_abundance" in uns. Counts + must have been generated with ``counts_from_abundance=None``. + (default: ``False``) + Attributes ---------- X @@ -225,6 +234,7 @@ def __init__( inference: Inference | None = None, quiet: bool = False, low_memory: bool = False, + from_pytximport: bool = False, ) -> None: # Initialize the AnnData part if adata is not None: @@ -318,6 +328,11 @@ def __init__( self.control_genes = control_genes self.logmeans = None self.filtered_genes = None + self.from_pytximport = from_pytximport + + if self.from_pytximport: + print("Detected pytximport data with length offsets.") + self._validate_pytximport_data() if inference: if n_cpus: @@ -645,6 +660,11 @@ def fit_size_factors( else: _control_mask = np.ones(self.X.shape[1], dtype=bool) + if fit_type not in [None, "ratio"] and self.from_pytximport: + raise ValueError( + "When from_pytximport=True, fit_type must be 'ratio' (default)." + ) + if fit_type == "iterative": self._fit_iterate_size_factors() @@ -682,8 +702,56 @@ def sizeFactor(x): UserWarning, stacklevel=2, ) + if self.from_pytximport: + raise ValueError( + "When from_pytximport=True, size factors cannot be computed " + "iteratively. Please check your input data." + ) + self._fit_iterate_size_factors() + elif self.from_pytximport: + if not self.quiet: + print( + "Using pytximport length offsets for normalization factors...", + file=sys.stderr, + ) + + # Create normalization matrix from length matrix + # Divide each gene's length by the geometric mean across samples + length_matrix = self.obsm["length"] + + with np.errstate(divide="ignore"): + # Gene-wise geometric means + log_geom_means = np.mean(np.log(length_matrix), axis=0) + norm_matrix = length_matrix / np.exp(log_geom_means) + + # Compute normalization factors using the DESeq2 algorithm + ( + counts_adj, + self.layers["normed_counts"], + norm_factors, + logmeans, + filtered_genes, + ) = estimate_norm_factors(self.X, norm_matrix, _control_mask) + + self.obsm["normalization_factors"] = norm_factors + self.logmeans = logmeans + self.filtered_genes = filtered_genes + + # Calculate the size factors on the normed counts + _, self.obs["size_factors"] = deseq2_norm_transform( + counts_adj, logmeans, filtered_genes + ) + + # Store sample-wise geometric mean of norm factors as size_factors + # This maintains compatibility with existing code while incorporating + # the gene-length correction + # with np.errstate(divide="ignore"): + # log_norm_factors = np.log(norm_factors) + # log_geom_mean_per_sample = np.mean(log_norm_factors, axis=1) + # self.obs["size_factors"] = np.exp(log_geom_mean_per_sample) + else: self.logmeans, self.filtered_genes = deseq2_norm_fit(self.X) _control_mask &= self.filtered_genes @@ -744,6 +812,12 @@ def fit_genewise_dispersions(self, vst=False) -> None: min_mu=self.min_mu, ) else: + # Use normalization factors if available + norm_factors = ( + self.obsm["normalization_factors"][:, self.non_zero_idx] + if "normalization_factors" in self.obsm + else None + ) _, mu_hat_, _, _ = self.inference.irls( counts=self.X[:, self.non_zero_idx], size_factors=size_factors, @@ -751,6 +825,7 @@ def fit_genewise_dispersions(self, vst=False) -> None: disp=self.var.loc[self.var["non_zero"], "_MoM_dispersions"].values, min_mu=self.min_mu, beta_tol=self.beta_tol, + normalization_factors=norm_factors, ) mu_param_name = "_vst_mu_hat" if vst else "_mu_hat" @@ -939,6 +1014,12 @@ def fit_LFC(self) -> None: if not self.quiet: print("Fitting LFCs...", file=sys.stderr) start = time.time() + # Use normalization factors if available + norm_factors = ( + self.obsm["normalization_factors"][:, self.non_zero_idx] + if "normalization_factors" in self.obsm + else None + ) mle_lfcs_, mu_, hat_diagonals_, converged_ = self.inference.irls( counts=self.X[:, self.non_zero_idx], size_factors=self.obs["size_factors"].values, @@ -946,6 +1027,7 @@ def fit_LFC(self) -> None: disp=self.var.loc[self.var["non_zero"], "dispersions"].values, min_mu=self.min_mu, beta_tol=self.beta_tol, + normalization_factors=norm_factors, ) end = time.time() @@ -1548,3 +1630,43 @@ def _check_full_rank_design(self): UserWarning, stacklevel=2, ) + + def _validate_pytximport_data(self): + """Validate that the data has the necessary pytximport information.""" + if "length" not in self.obsm_keys(): + raise ValueError( + "from_pytximport=True but 'length' not found in obsm. " + "Make sure your AnnData object was created from pytximport output." + ) + + if "counts_from_abundance" not in self.uns_keys(): + raise ValueError( + "from_pytximport=True but 'counts_from_abundance' not found in uns. " + "Make sure your AnnData object was created from pytximport output." + ) + + # Check that counts_from_abundance is None (meaning raw counts were used) + if self.uns["counts_from_abundance"] is not None: + raise ValueError( + "Normalization factors can only be used with raw counts. " + f"Got counts_from_abundance={self.uns['counts_from_abundance']}. " + "Please use pytximport with counts_from_abundance=None." + ) + + # Validate length matrix dimensions + length_matrix = self.obsm["length"] + if length_matrix.shape != (self.n_obs, self.n_vars): + raise ValueError( + f"Length matrix shape {length_matrix.shape} does not match " + f"count matrix shape {(self.n_obs, self.n_vars)}" + ) + + # Check for non-positive lengths + if np.any(length_matrix <= 0): + raise ValueError( + "Length matrix contains non-positive values. " + "All gene lengths must be positive." + ) + + if not self.quiet: + print("Detected pytximport data with length offsets.", file=sys.stderr) diff --git a/pydeseq2/default_inference.py b/pydeseq2/default_inference.py index 9bd67bad..ed3733ca 100644 --- a/pydeseq2/default_inference.py +++ b/pydeseq2/default_inference.py @@ -92,6 +92,7 @@ def irls( # noqa: D102 max_beta: float = 30, optimizer: Literal["BFGS", "L-BFGS-B"] = "L-BFGS-B", maxiter: int = 250, + normalization_factors: np.ndarray | None = None, ) -> tuple[np.ndarray, np.ndarray, np.ndarray, np.ndarray]: with parallel_backend(self._backend, inner_max_num_threads=1): res = Parallel( @@ -110,6 +111,11 @@ def irls( # noqa: D102 max_beta=max_beta, optimizer=optimizer, maxiter=maxiter, + normalization_factors=( + normalization_factors[:, i] + if normalization_factors is not None + else None + ), ) for i in range(counts.shape[1]) ) diff --git a/pydeseq2/ds.py b/pydeseq2/ds.py index 3419607f..03780f42 100644 --- a/pydeseq2/ds.py +++ b/pydeseq2/ds.py @@ -316,11 +316,20 @@ def run_wald_test(self) -> None: file=sys.stderr, ) - mu = ( - np.exp(self.design_matrix @ self.LFC.T) - .multiply(self.dds.obs["size_factors"], 0) - .values - ) + # Use normalization factors if available, otherwise use size factors + if "normalization_factors" in self.dds.obsm: + # For normalization factors: mu = exp(X @ beta) * norm_factors + # Both matrices are (samples, genes) + exp_design_lfc = np.exp(self.design_matrix @ self.LFC.T).values + mu = exp_design_lfc * self.dds.obsm["normalization_factors"] + else: + # Standard case: mu = exp(X @ beta) * size_factors + # Result is (samples, genes), size_factors is (samples,) + mu = ( + np.exp(self.design_matrix @ self.LFC.T) + .multiply(self.dds.obs["size_factors"], 0) + .values + ) # Set regularization factors. if self.prior_LFC_var is not None: @@ -384,7 +393,11 @@ def lfc_shrink(self, coeff: str, adapt: bool = True) -> None: design_matrix = self.design_matrix.values size = 1.0 / self.dds.var["dispersions"].values - offset = np.log(self.dds.obs["size_factors"]).values + + if "normalization_factors" in self.dds.obsm: + offset = np.log(self.dds.obsm["normalization_factors"]) + else: + offset = np.log(self.dds.obs["size_factors"].values) # Set priors prior_no_shrink_scale = 15 diff --git a/pydeseq2/inference.py b/pydeseq2/inference.py index 40bff73b..e79085fc 100644 --- a/pydeseq2/inference.py +++ b/pydeseq2/inference.py @@ -55,6 +55,7 @@ def irls( max_beta: float = 30, optimizer: Literal["BFGS", "L-BFGS-B"] = "L-BFGS-B", maxiter: int = 250, + normalization_factors: np.ndarray | None = None, ) -> tuple[np.ndarray, np.ndarray, np.ndarray, np.ndarray]: r"""Fit a NB GLM wit log-link to predict counts from the design matrix. diff --git a/pydeseq2/preprocessing.py b/pydeseq2/preprocessing.py index 0c0a5a34..d2dc2cac 100644 --- a/pydeseq2/preprocessing.py +++ b/pydeseq2/preprocessing.py @@ -100,3 +100,82 @@ def deseq2_norm_transform( size_factors = np.exp(log_medians) deseq2_counts = counts / size_factors[:, None] return deseq2_counts, size_factors + + +def estimate_norm_factors( + counts: pd.DataFrame | np.ndarray, + norm_matrix: np.ndarray, + control_mask: np.ndarray | None = None, +) -> tuple[ + pd.DataFrame | np.ndarray, + pd.DataFrame | np.ndarray, + np.ndarray, + np.ndarray, + np.ndarray, +]: + """Estimate normalization factors from counts and normalization matrix. + + Implements the DESeq2 estimateNormFactors algorithm for pytximport integration. + This function computes normalization factors that account for both library size + and gene length differences. + + Parameters + ---------- + counts : pandas.DataFrame or ndarray + Raw counts. One column per gene, one row per sample. + + norm_matrix : ndarray + Normalization matrix. Should have the same dimensions as counts. + + control_mask : ndarray, optional + Boolean mask indicating which genes to use as control genes for size + factor estimation. If None, all genes are used. + + Returns + ------- + counts_adj : pandas.DataFrame or ndarray + Counts adjusted by the normalization matrix. + + normed_counts : pandas.DataFrame or ndarray + Normalized counts using the computed normalization factors. + + norm_factors : ndarray + Computed normalization factors that account for both library size + and gene length differences. + + logmeans : ndarray + Gene-wise mean log counts from the adjusted counts. + + filtered_genes : ndarray + Genes whose log means are different from -∞ in the adjusted counts. + """ + # Divide counts by normalization matrix + counts_adj = counts / norm_matrix + + # Compute size factors from adjusted counts using standard DESeq2 method + logmeans, filtered_genes = deseq2_norm_fit(counts_adj) + + # Apply control gene mask if provided + if control_mask is not None: + filtered_genes = filtered_genes & control_mask + + _, size_factors = deseq2_norm_transform(counts_adj, logmeans, filtered_genes) + + # Compute normalization factors: (norm_matrix.T * size_factors).T + # Broadcasting: norm_matrix (samples x genes) * size_factors (samples,) + norm_factors = norm_matrix * size_factors[:, None] + + # Normalize to geometric mean of 1 (row-wise geometric mean) + with np.errstate(divide="ignore"): # ignore division by zero warnings + log_norm_factors = np.log(norm_factors) + log_geom_means = np.mean(log_norm_factors, axis=1, keepdims=True) + norm_factors = norm_factors / np.exp(log_geom_means) + + # Compute normalized counts + normed_counts = counts / norm_factors + + # TODO: Check whether we actually have to calculate logmeans and filtered_genes again + # from the normed counts, or if we can just return the previous ones. + logmeans, filtered_genes = deseq2_norm_fit(normed_counts) + + return counts_adj, normed_counts, norm_factors, logmeans, filtered_genes diff --git a/pydeseq2/utils.py b/pydeseq2/utils.py index c538c137..7aa20382 100644 --- a/pydeseq2/utils.py +++ b/pydeseq2/utils.py @@ -283,6 +283,7 @@ def irls_solver( max_beta: float = 30, optimizer: Literal["BFGS", "L-BFGS-B"] = "L-BFGS-B", maxiter: int = 250, + normalization_factors: np.ndarray | None = None, ) -> tuple[np.ndarray, np.ndarray, np.ndarray, bool]: r"""Fit a NB GLM wit log-link to predict counts from the design matrix. @@ -327,6 +328,11 @@ def irls_solver( Maximum number of IRLS iterations to perform before switching to L-BFGS-B. (default: ``250``). + normalization_factors : ndarray, optional + Gene-specific normalization factors (samples x genes). If provided, these + will be used instead of size_factors for computing mu. This enables support + for pytximport length-corrected normalization. (default: ``None``). + Returns ------- beta: ndarray @@ -347,28 +353,36 @@ def irls_solver( num_vars = design_matrix.shape[1] X = design_matrix + # Choose appropriate normalization factors + if normalization_factors is not None: + # Use gene-specific normalization factors (1D array for single gene) + norm_factors = normalization_factors + else: + # Use standard size factors (1D array) + norm_factors = size_factors + # if full rank, estimate initial betas for IRLS below if np.linalg.matrix_rank(X) == num_vars: Q, R = np.linalg.qr(X) - y = np.log(counts / size_factors + 0.1) + y = np.log(counts / norm_factors + 0.1) beta_init = solve(R, Q.T @ y) beta = beta_init else: # Initialise intercept with log base mean beta_init = np.zeros(num_vars) - beta_init[0] = np.log(counts / size_factors).mean() + beta_init[0] = np.log(counts / norm_factors).mean() beta = beta_init dev = 1000.0 dev_ratio = 1.0 ridge_factor = np.diag(np.repeat(1e-6, num_vars)) - mu = np.maximum(size_factors * np.exp(X @ beta), min_mu) + mu = np.maximum(norm_factors * np.exp(X @ beta), min_mu) converged = True i = 0 while dev_ratio > beta_tol: W = mu / (1.0 + mu * disp) - z = np.log(mu / size_factors) + (counts - mu) / mu + z = np.log(mu / norm_factors) + (counts - mu) / mu H = (X.T * W) @ X + ridge_factor beta_hat = solve(H, X.T @ (W * z), assume_a="pos") i += 1 @@ -377,11 +391,11 @@ def irls_solver( # If IRLS starts diverging, use L-BFGS-B def f(beta: np.ndarray) -> float: # closure to minimize - mu_ = np.maximum(size_factors * np.exp(X @ beta), min_mu) + mu_ = np.maximum(norm_factors * np.exp(X @ beta), min_mu) return nb_nll(counts, mu_, disp) + 0.5 * (ridge_factor @ beta**2).sum() def df(beta: np.ndarray) -> np.ndarray: - mu_ = np.maximum(size_factors * np.exp(X @ beta), min_mu) + mu_ = np.maximum(norm_factors * np.exp(X @ beta), min_mu) return ( -X.T @ counts + ((1 / disp + counts) * mu_ / (1 / disp + mu_)) @ X @@ -401,7 +415,7 @@ def df(beta: np.ndarray) -> np.ndarray: ) beta = res.x - mu = np.maximum(size_factors * np.exp(X @ beta), min_mu) + mu = np.maximum(norm_factors * np.exp(X @ beta), min_mu) converged = res.success if not res.success and num_vars <= 2: @@ -411,11 +425,11 @@ def df(beta: np.ndarray) -> np.ndarray: X, disp, ) - mu = np.maximum(size_factors * np.exp(X @ beta), min_mu) + mu = np.maximum(norm_factors * np.exp(X @ beta), min_mu) break beta = beta_hat - mu = np.maximum(size_factors * np.exp(X @ beta), min_mu) + mu = np.maximum(norm_factors * np.exp(X @ beta), min_mu) # Compute deviation old_dev = dev # Replaced deviation with -2 * nll, as in the R code @@ -436,7 +450,7 @@ def df(beta: np.ndarray) -> np.ndarray: # Return an UNthresholded mu (as in the R code) # Previous quantities are estimated with a threshold though - mu = size_factors * np.exp(X @ beta) + mu = norm_factors * np.exp(X @ beta) return beta, mu, H, converged diff --git a/pyproject.toml b/pyproject.toml index eecde408..ef43aa22 100644 --- a/pyproject.toml +++ b/pyproject.toml @@ -3,6 +3,8 @@ line-length = 89 [tool.ruff] line-length = 89 + +[tool.ruff.lint] select = [ "F", # Errors detected by Pyflakes "E", # Error detected by Pycodestyle @@ -29,13 +31,13 @@ ignore = [ "D213", ] -[tool.ruff.isort] +[tool.ruff.lint.isort] force-single-line = true -[tool.ruff.pydocstyle] +[tool.ruff.lint.pydocstyle] convention = "numpy" -[tool.ruff.per-file-ignores] +[tool.ruff.lint.per-file-ignores] "docs/*" = ["I"] "tests/*" = ["D"] # Ignore errors linked to sphinx gallery syntax diff --git a/tests/data/pytximport/test_counts.csv b/tests/data/pytximport/test_counts.csv new file mode 100644 index 00000000..099b7e7d --- /dev/null +++ b/tests/data/pytximport/test_counts.csv @@ -0,0 +1,41128 @@ +,SRR16504309_wt,SRR16504310_wt,SRR16504311_ko,SRR16504312_ko +DDX11L16,10.223884781831023,0.0,8.509129631093488,8.900706496998497 +DDX11L1,0.0,0.0,0.0,0.0 +WASH7P,9.723257170323963,0.0,18.100530363393283,22.038945887849746 +MIR6859-1,0.0,0.0,0.0,0.0 +MIR1302-2HG,0.0,0.0,0.0,0.0 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+MT-ND6,10252.561934929592,5352.202862161514,3778.200383952726,4014.363563297372 +MT-TE,281.83998660841434,141.90429680944882,95.7825881214228,88.24932112769015 +MT-CYB,20781.660560327375,10496.983366251246,10651.193870372417,10737.00052943413 +MT-TT,62.08363341374163,38.16736259012762,16.873616560250646,17.15959021927309 +MT-TP,88.69090487677376,45.9965651727179,28.784404720427574,23.533152300717376 diff --git a/tests/data/pytximport/test_metadata.csv b/tests/data/pytximport/test_metadata.csv new file mode 100644 index 00000000..e1da0301 --- /dev/null +++ b/tests/data/pytximport/test_metadata.csv @@ -0,0 +1,5 @@ +,condition +SRR16504309_wt,wt +SRR16504310_wt,wt +SRR16504311_ko,ko +SRR16504312_ko,ko diff --git a/tests/data/pytximport/test_pytximport.h5ad b/tests/data/pytximport/test_pytximport.h5ad new file mode 100644 index 00000000..932fb725 Binary files /dev/null and b/tests/data/pytximport/test_pytximport.h5ad differ diff --git a/tests/data/pytximport/test_pytximport_length_scaled.h5ad b/tests/data/pytximport/test_pytximport_length_scaled.h5ad new file mode 100644 index 00000000..0255d74c Binary files /dev/null and b/tests/data/pytximport/test_pytximport_length_scaled.h5ad differ diff --git a/tests/test_pytximport.py b/tests/test_pytximport.py new file mode 100644 index 00000000..6696fe98 --- /dev/null +++ b/tests/test_pytximport.py @@ -0,0 +1,240 @@ +import os +from pathlib import Path + +import anndata as ad +import numpy as np +import pandas as pd +import pytest + +import tests +from pydeseq2.dds import DeseqDataSet +from pydeseq2.ds import DeseqStats + + +@pytest.fixture +def pytximport_adata(): + """Load test pytximport AnnData object.""" + test_path = str(Path(os.path.realpath(tests.__file__)).parent.resolve()) + adata_path = os.path.join(test_path, "data/pytximport/test_pytximport.h5ad") + return ad.read_h5ad(adata_path) + + +@pytest.fixture +def standard_adata(): + """Create standard AnnData object without pytximport fields.""" + test_path = str(Path(os.path.realpath(tests.__file__)).parent.resolve()) + + counts_path = os.path.join(test_path, "data/pytximport/test_counts.csv") + metadata_path = os.path.join(test_path, "data/pytximport/test_metadata.csv") + + counts_df = pd.read_csv(counts_path, index_col=0) + metadata_df = pd.read_csv(metadata_path, index_col=0) + + adata = ad.AnnData(X=counts_df.T.values.round(), obs=metadata_df) + adata.var_names = counts_df.index + adata.obs_names = counts_df.columns + + return adata + + +class TestPytximportDetection: + """Test pytximport data detection and validation.""" + + def test_explicit_flag(self, pytximport_adata): + """Test explicit from_pytximport=True flag.""" + dds = DeseqDataSet( + adata=pytximport_adata, design="~condition", from_pytximport=True + ) + + # Should have detected pytximport data during initialization + assert dds.from_pytximport is True + + dds.fit_size_factors() + assert "normalization_factors" in dds.obsm + assert dds.obsm["normalization_factors"].shape == (4, 41127) + + def test_no_pytximport_data(self, standard_adata): + """Test standard DESeq2 without pytximport data.""" + dds = DeseqDataSet(adata=standard_adata, design="~condition") + + dds.fit_size_factors() + assert "normalization_factors" not in dds.obsm + assert "size_factors" in dds.obs + + +class TestPytximportValidation: + """Test pytximport data validation.""" + + def test_missing_length_matrix(self, standard_adata): + """Test validation when length matrix is missing.""" + with pytest.raises(ValueError, match="'length' not found in obsm"): + DeseqDataSet(adata=standard_adata, design="~condition", from_pytximport=True) + + def test_missing_counts_from_abundance(self, pytximport_adata): + """Test validation when counts_from_abundance is missing.""" + # Remove the counts_from_abundance key + del pytximport_adata.uns["counts_from_abundance"] + + with pytest.raises(ValueError, match="'counts_from_abundance' not found in uns"): + DeseqDataSet( + adata=pytximport_adata, design="~condition", from_pytximport=True + ) + + def test_invalid_counts_from_abundance(self, pytximport_adata): + """Test validation when counts are not raw counts.""" + pytximport_adata.uns["counts_from_abundance"] = "scaledTPM" + + with pytest.raises(ValueError, match="only be used with raw counts"): + DeseqDataSet( + adata=pytximport_adata, design="~condition", from_pytximport=True + ) + + def test_length_matrix_shape(self, pytximport_adata): + """Test validation of length matrix shape.""" + # Create a copy to avoid modifying the fixture + test_adata = pytximport_adata.copy() + + # Test wrong number of columns (genes) - this should pass AnnData validation + # but fail our pytximport validation + test_adata.obsm["length"] = np.random.rand(4, 30) # Wrong n_vars + + with pytest.raises(ValueError, match="Length matrix shape"): + DeseqDataSet(adata=test_adata, design="~condition", from_pytximport=True) + + def test_negative_lengths(self, pytximport_adata): + """Test validation of non-positive gene lengths.""" + # Add some negative lengths + length_matrix = pytximport_adata.obsm["length"].copy() + length_matrix[0, 0] = -1.0 + pytximport_adata.obsm["length"] = length_matrix + + with pytest.raises(ValueError, match="non-positive values"): + DeseqDataSet( + adata=pytximport_adata, design="~condition", from_pytximport=True + ) + + +class TestNormalizationFactors: + """Test normalization factor computation and usage.""" + + def test_normalization_factor_computation(self, pytximport_adata): + """Test that normalization factors are computed correctly.""" + dds = DeseqDataSet( + adata=pytximport_adata, + design="~condition", + from_pytximport=True, + ) + dds.fit_size_factors() + + # Check normalization factors properties + norm_factors = dds.obsm["normalization_factors"] + assert norm_factors.shape == (4, 41127) + assert np.all(norm_factors > 0) + + # Check that they're different from simple size factors + # (due to gene-length correction) + size_factors = dds.obs["size_factors"].values + norm_factor_means = norm_factors.mean(axis=1) + + # Check that normalization factors vary across genes (not constant) + norm_factor_std = np.std(norm_factors, axis=1) + assert np.all(norm_factor_std > 0) # Should have variation across genes + + # Should have some difference from size factors + max_diff = np.max(np.abs(norm_factor_means - size_factors)) + assert max_diff > 0.001 # At least some difference + + def test_normalized_counts(self, pytximport_adata): + """Test that normalized counts are computed correctly.""" + dds = DeseqDataSet( + adata=pytximport_adata, + design="~condition", + from_pytximport=True, + ) + dds.fit_size_factors() + + # Check normalized counts + normed_counts = dds.layers["normed_counts"] + assert normed_counts.shape == (4, 41127) + assert np.all(normed_counts >= 0) + + # Normalized counts should be different from simple division + simple_normed = dds.X / dds.obs["size_factors"].values[:, None] + assert not np.allclose(normed_counts, simple_normed, rtol=0.01) + + def test_comparison_with_standard_deseq2(self, pytximport_adata, standard_adata): + """Compare pytximport and standard DESeq2 normalization.""" + # pytximport normalization + dds_pyxtm = DeseqDataSet( + adata=pytximport_adata, + design="~condition", + from_pytximport=True, + ) + dds_pyxtm.fit_size_factors() + + # Standard normalization + dds_std = DeseqDataSet(adata=standard_adata, design="~condition") + dds_std.fit_size_factors() + + # The main difference should be in having normalization factors vs not + assert "normalization_factors" in dds_pyxtm.obsm + assert "normalization_factors" not in dds_std.obsm + + # Both should have size factors + assert "size_factors" in dds_pyxtm.obs + assert "size_factors" in dds_std.obs + + # Normalization factors should exist and have correct properties + norm_factors = dds_pyxtm.obsm["normalization_factors"] + assert norm_factors.shape == (4, 41127) + assert np.all(norm_factors > 0) + + +class TestDownstreamAnalysis: + """Test that downstream analysis works with normalization factors.""" + + def test_deseq2_pipeline_with_normalization_factors(self, pytximport_adata): + """Test full DESeq2 pipeline with pytximport normalization factors.""" + dds = DeseqDataSet( + adata=pytximport_adata, design="~condition", from_pytximport=True + ) + + # Run full DESeq2 pipeline - may have convergence warnings with small data + with pytest.warns( + UserWarning, match="As the residual degrees of freedom is less than 3" + ): + dds.deseq2() + + # Check that all expected results are computed + assert "LFC" in dds.varm + assert "dispersions" in dds.var + assert "size_factors" in dds.obs + assert "normalization_factors" in dds.obsm + + # Test differential expression analysis + stat_res = DeseqStats(dds, contrast=["condition", "ko", "wt"]) + stat_res.summary() + + # Check results + assert hasattr(stat_res, "results_df") + assert "log2FoldChange" in stat_res.results_df.columns + assert "padj" in stat_res.results_df.columns + + # Results should be reasonable + results_df = stat_res.results_df + assert results_df.shape[0] == 41127 # All genes + assert not results_df["log2FoldChange"].isna().all() + + def test_vst_with_normalization_factors(self, pytximport_adata): + """Test VST transformation with pytximport normalization factors.""" + dds = DeseqDataSet( + adata=pytximport_adata, design="~condition", from_pytximport=True + ) + + dds.vst() + + # Check VST results + assert "vst_counts" in dds.layers + vst_counts = dds.layers["vst_counts"] + assert vst_counts.shape == (4, 41127) + assert np.all(np.isfinite(vst_counts))