Introduction

Cell-type labels from different datasets are often inconsistent. Authors use different naming conventions, levels of granularity, and terminology. For example:

  • “Purkinje neurons” vs “Purkinje cells” vs “PCs”
  • “Astrocytes” vs “Astrocyte-like cells” vs “GFAP+ glia”
  • “Inhibitory neurons” vs “GABAergic neurons” vs “Interneurons”

scNLP provides NLP-based approaches to identify semantically similar labels and harmonize cell-type annotations across datasets without requiring manual curation.

Example Data

We’ll use the included pseudo-bulk dataset containing samples from multiple studies with heterogeneous cell-type labels:

data("pseudo_seurat")

# View the diversity of labels and sources
label_summary <- table(
    Dataset = pseudo_seurat$batch,
    Species = pseudo_seurat$species
)
print(label_summary)
##                                 Species
## Dataset                          fly human mouse zebrafish
##   AIBS                             0     6     0         0
##   Aerts2020                      116     0     0         0
##   BlueLake2018_FrontalCortexOnly   0    24     0         0
##   BlueLake2018_VisualCortexOnly    0    28     0         0
##   DRONC_human                      0    15     0         0
##   DRONC_mouse                      0     0    22         0
##   LaManno2020                      0     0   206         0
##   Raj2020                          0     0     0       222
##   Zeisel2016                       0     0    48         0
##   Zeisel2018                       0     0    37         0
##   descartes_SampledData            0    77     0         0
# Count unique cell-type labels
length(unique(pseudo_seurat$celltype))
## [1] 608
# Sample of label heterogeneity
head(unique(pseudo_seurat$celltype), 20)
##  [1] ASC1   ASC2   END    exCA1  exCA3  exDG   exPFC1 exPFC2 GABA1  GABA2 
## [11] MG     NSC    ODC1   ODC2   OPC    ChP    exPFC3 exPFC4 exPFC5 exPFC6
## 608 Levels: A AFP_ALB_positive_cells ASC1 ASC2 ... ventral_midbrain

Strategy 1: TF-IDF Clustering

TF-IDF analysis identifies enriched terms within each cluster. Cells with similar enriched terms likely represent the same cell type, regardless of the exact label used.

Compute TF-IDF

# Run TF-IDF on the existing clusters
result <- run_tfidf(
    obj = pseudo_seurat,
    reduction = "umap",
    cluster_var = "cluster",
    label_var = "celltype"
)
## Extracting obsm from Seurat: umap
## + Dropping 2 conflicting obs variables: UMAP.1, UMAP.2
## Loading required namespace: tidytext
## Setting cell metadata (obs) in obj.

Visualize Enriched Terms

The scatter plot shows cells colored by cluster, with enriched terms labeled at cluster centers:

res <- plot_tfidf(
    obj = pseudo_seurat,
    label_var = "celltype",
    cluster_var = "cluster",
    show_plot = TRUE
)
## Extracting obsm from Seurat: umap
## + Dropping 2 conflicting obs variables: UMAP.1, UMAP.2
## Setting cell metadata (obs) in obj.
## Warning: `aes_string()` was deprecated in ggplot2 3.0.0.
##  Please use tidy evaluation idioms with `aes()`.
##  See also `vignette("ggplot2-in-packages")` for more information.
##  The deprecated feature was likely used in the scNLP package.
##   Please report the issue at <https://github.com/neurogenomics/scNLP/issues>.
## This warning is displayed once per session.
## Call `lifecycle::last_lifecycle_warnings()` to see where this warning was
## generated.
## Warning in ggplot2::geom_point(ggplot2::aes_string(color = color_var, size =
## size_var, : Ignoring unknown aesthetics: label
## Warning: Using `size` aesthetic for lines was deprecated in ggplot2 3.4.0.
##  Please use `linewidth` instead.
##  The deprecated feature was likely used in the scNLP package.
##   Please report the issue at <https://github.com/neurogenomics/scNLP/issues>.
## This warning is displayed once per session.
## Call `lifecycle::last_lifecycle_warnings()` to see where this warning was
## generated.

Examine TF-IDF Results

The TF-IDF results reveal which terms are most distinctive for each cluster:

# Get the full TF-IDF results
tfidf_df <- res$tfidf_df

# Top terms per cluster
top_terms <- tfidf_df |>
    dplyr::group_by(cluster) |>
    dplyr::slice_max(order_by = tf_idf, n = 3) |>
    dplyr::select(cluster, word, tf_idf)

print(top_terms, n = 30)
## # A tibble: 44 × 3
## # Groups:   cluster [15]
##    cluster word       tf_idf
##    <fct>   <chr>       <dbl>
##  1 0       lpn        0.0528
##  2 0       adpn       0.0523
##  3 0       neuron     0.0428
##  4 1       rhombomere 0.154 
##  5 1       placode    0.116 
##  6 1       hindbrain  0.114 
##  7 2       neurons    0.101 
##  8 2       glia       0.0587
##  9 2       neuron     0.0419
## 10 3       fibroblast 0.249 
## 11 3       arch       0.0653
## 12 4       e7         0.112 
## 13 4       e8         0.0612
## 14 4       neural     0.0596
## 15 5       glia       0.198 
## 16 5       schwann    0.182 
## 17 5       radial     0.112 
## 18 6       neurons    0.239 
## 19 6       peripheral 0.104 
## 20 6       neuron     0.0704
## 21 7       gaba1      0.120 
## 22 7       gaba2      0.120 
## 23 7       in1a       0.120 
## 24 8       opc        0.143 
## 25 8       radial     0.119 
## 26 8       opcs       0.102 
## 27 9       vascular   0.528 
## 28 9       peric      0.0423
## 29 9       pericytes  0.0423
## 30 10      immune     0.521 
## # ℹ 14 more rows

Clusters sharing similar enriched terms (e.g., “neuron”, “astrocyte”, “oligodendrocyte”) likely contain the same cell types from different datasets.

Strategy 2: Neighbor-based Harmonization

The search_neighbors() function finds cells with similar expression profiles, allowing you to identify which labels from different datasets correspond to the same cell types.

Find Similar Cells

Search for cells similar to those labeled “purkinje”:

purkinje_neighbors <- search_neighbors(
    seurat = pseudo_seurat,
    var1_search = "purkinje",
    max_neighbors = 10
)
## No variable features detected. Computing
## No PCA detected. Computing
## Centering and scaling data matrix
## PC_ 1 
## Positive:  FTMT, ACTG1, ALAS2, HSPA1L, NME1-NME2, POTEI, OTOP2, RAPSN, BEST2, DPEP2 
##     HMGB2, NAPRT1, KRT8, PPIB, DSCC1, POU4F3, CCDC102A, GAPDHS, CHST6, AGXT2 
##     LYZL2, MTMR8, ACTG2, ACPL2, BANF1, PPAPDC2, HTR1A, IFI30, CYBRD1, LHX8 
## Negative:  FAIM2, CAMK2A, SCN1A, CAMK2B, FRRS1L, UNC80, PHYHIP, RASGRF2, CCK, GRIA2 
##     STXBP5L, ARPP21, SLC12A5, DIRAS2, RYR2, SLC4A10, KCNT1, GRM5, CAMKV, KIAA1211L 
##     GABRA4, GABRA1, SV2B, CX3CL1, AK5, PNMA2, JPH4, DGKG, GPR158, KCNC2 
## PC_ 2 
## Positive:  CAMKK1, DGKQ, NT5DC3, CA7, ABCG4, HTR1A, C5orf28, OTOP2, HYKK, DPEP2 
##     CHST6, POTEI, SLC8A3, SLC38A11, ADRA2A, MPPED1, MTMR8, HTR7, CACNA1B, PPAPDC2 
##     C2orf69, GRIK1, IFI30, STK32B, RASL10B, SLC24A4, FAXDC2, ADCY3, ACSS2, ANKRD29 
## Negative:  RAN, HSP90AA1, H2AFZ, HNRNPAB, CCT5, NPM1, GNG5, DBI, HMGB2, ITM2B 
##     ATP6V1G1, SERPINH1, CIRBP, CD63, NDUFA6, MDK, JUN, MYL12B, SPARC, NPC2 
##     GLUL, ID3, EEF1A1, VIM, CLIC1, COX6B1, LDHA, DDAH2, ENO1, CNN3 
## PC_ 3 
## Positive:  ADGRL2, AC011288.2, RP11-420N3.3, RP11-191L9.4, NRXN3, PLPPR1, RP11-123O10.4, ZNF385D, AC114765.1, NWD2 
##     RBFOX3, MIR137HG, MIR325HG, SGOL1-AS1, POU6F2, ANKRD18A, LY86-AS1, LINC01197, DGCR5, DPY19L1P1 
##     MIR4300HG, AQP4-AS1, HPSE2, LINC00632, NLGN4X, AC067956.1, PWRN1, LINC00599, CABP1, LINC01158 
## Negative:  KRTCAP2, APOE, C20orf24, PDIA6, PGLS, GNG11, S100A13, HIST1H2BI, ISCA2, GSTM5 
##     LAPTM4A, CST3, TMEM176B, KLF4, PDLIM2, CAP1, S100A16, APRT, CYR61, FAIM 
##     IFITM3, CDKN1A, KLF2, CLIC1, ARPC1B, IER2, S100A1, CMTM5, FXYD1, TCN2 
## PC_ 4 
## Positive:  RESP18, CTXN2, ATP6V1G2, GNG13, DISP2, C15orf59, CCDC85A, GNG3, SYNGR3, RGS8 
##     VWA5B2, C1QL3, HPCA, TUBB3, CALB1, SNCB, HTR3A, ARHGDIG, L1CAM, NAP1L5 
##     PCDH20, HMP19, DBNDD2, NPAS4, FABP3, CALY, FAM43B, CKMT1B, LOC728392, LTK 
## Negative:  PTPN18, SLCO1A2, LINC00639, INPP5D, IFI44, LYN, DISC1, NEAT1, NRGN, CMYA5 
##     IFI44L, GALNT15, PARP14, AC012593.1, AQP4-AS1, MSR1, MT2A, ISG15, SHROOM4, CABP1 
##     UACA, KCNQ1OT1, PART1, CNDP1, FAM153B, DGCR5, SOX2-OT, LINC00844, ADGRG1, LINC00599 
## PC_ 5 
## Positive:  MEST, IGFBP2, CNN3, FBXL7, NNAT, TUBB2B, GPC3, VIM, NKAIN4, ID1 
##     BMP7, CSRP2, NDN, DDAH2, GPX8, IGFBPL1, MARCKSL1, GSTM3, FBLN1, PARD3 
##     MFAP4, PTN, FABP7, COPS6, CTNNA2, ZBTB20, BEX1, CD81, ENO1, NPAS3 
## Negative:  C1QB, FCGR2A, MS4A6A, TYROBP, C1QC, AIF1, C1QA, CSF1R, CD86, MRC1 
##     MS4A7, CTSS, CCL24, FCER1G, CD53, CD14, FCGR1A, PLEK, C3AR1, LYZ 
##     FCGR2B, CX3CR1, CCL3L3, CCL2, CCR1, CD68, C5AR1, PF4, HPGDS, LY86
## No graphs detected. Computing.
## Computing nearest neighbor graph
## Computing SNN
## Using graph: RNA_snn
## + Filtering results by `var1_search`: purkinje
## + 3 entries matching `var1_search` identified.
## + Adding original names to results
## + Returning 32 pair-wise similarities.
head(purkinje_neighbors)
##                                            Var1
##                                          <char>
## 1: human.descartes_SampledData.Purkinje_neurons
## 2: human.descartes_SampledData.Purkinje_neurons
## 3: human.descartes_SampledData.Purkinje_neurons
## 4: human.descartes_SampledData.Purkinje_neurons
## 5:           zebrafish.Raj2020.purkinje_neurons
## 6:           zebrafish.Raj2020.purkinje_neurons
##                                                      Var2 similarity
##                                                    <char>      <num>
## 1: human.descartes_SampledData.SATB2_LRRC7_positive_cells  0.9047619
## 2:           human.descartes_SampledData.Oligodendrocytes  0.8181818
## 3:            human.descartes_SampledData.Granule_neurons  0.8181818
## 4:                human.descartes_SampledData.ENS_neurons  0.8181818
## 5:    zebrafish.Raj2020.neurons._gabaergic._glutamatergic  0.8181818
## 6:      zebrafish.Raj2020.purkinje_neurons_.gabaergic_...  0.8181818
##                                         Var1_id
##                                          <char>
## 1: human.descartes_SampledData.Purkinje_neurons
## 2: human.descartes_SampledData.Purkinje_neurons
## 3: human.descartes_SampledData.Purkinje_neurons
## 4: human.descartes_SampledData.Purkinje_neurons
## 5:           zebrafish.Raj2020.purkinje_neurons
## 6:           zebrafish.Raj2020.purkinje_neurons
##                                                   Var2_id
##                                                    <char>
## 1: human.descartes_SampledData.SATB2_LRRC7_positive_cells
## 2:           human.descartes_SampledData.Oligodendrocytes
## 3:            human.descartes_SampledData.Granule_neurons
## 4:                human.descartes_SampledData.ENS_neurons
## 5:    zebrafish.Raj2020.neurons._gabaergic._glutamatergic
## 6:      zebrafish.Raj2020.purkinje_neurons_.gabaergic_...

Cross-dataset Mapping

Examine which labels from other datasets map to Purkinje cells:

# Get unique neighbor labels
neighbor_labels <- unique(purkinje_neighbors$Var2)
cat("Labels similar to 'purkinje':\n")
## Labels similar to 'purkinje':
print(neighbor_labels)
##  [1] "human.descartes_SampledData.SATB2_LRRC7_positive_cells"                 
##  [2] "human.descartes_SampledData.Oligodendrocytes"                           
##  [3] "human.descartes_SampledData.Granule_neurons"                            
##  [4] "human.descartes_SampledData.ENS_neurons"                                
##  [5] "zebrafish.Raj2020.neurons._gabaergic._glutamatergic"                    
##  [6] "zebrafish.Raj2020.purkinje_neurons_.gabaergic_..."                      
##  [7] "zebrafish.Raj2020.progenitors.differentiating_granule_cells_.hindbrain."
##  [8] "zebrafish.Raj2020.ventral_habenula"                                     
##  [9] "zebrafish.Raj2020.purkinje_neurons"                                     
## [10] "human.descartes_SampledData.Inhibitory_interneurons"                    
## [11] "human.descartes_SampledData.Excitatory_neurons"                         
## [12] "zebrafish.Raj2020.telencephalon_.pallium._neuron._glutamatergic"        
## [13] "zebrafish.Raj2020.cranial_ganglion"                                     
## [14] "zebrafish.Raj2020.neurons_.glutamatergic._midbrain."                    
## [15] "zebrafish.Raj2020.diencephalon.mid"                                     
## [16] "zebrafish.Raj2020.neurons_.gabaergic_..._midbrain._optic_tectum."       
## [17] "human.descartes_SampledData.SKOR2_NPSR1_positive_cells"                 
## [18] "zebrafish.Raj2020.retina_neurons_.horizontal.."                         
## [19] "zebrafish.Raj2020.neurons_.no_neurod1.neurod4._glutamatergic"           
## [20] "human.descartes_SampledData.SLC24A4_PEX5L_positive_cells"               
## [21] "human.descartes_SampledData.Inhibitory_neurons"                         
## [22] "human.descartes_SampledData.Amacrine_cells"                             
## [23] "human.descartes_SampledData.PDE11A_FAM19A2_positive_cells"

Search Multiple Cell Types

You can search for multiple cell types to build a harmonization map:

# Search for astrocyte-like cells
astro_neighbors <- search_neighbors(
    seurat = pseudo_seurat,
    var1_search = "astro",
    max_neighbors = 5
)
## No variable features detected. Computing
## No PCA detected. Computing
## Centering and scaling data matrix
## PC_ 1 
## Positive:  FTMT, ACTG1, ALAS2, HSPA1L, NME1-NME2, POTEI, OTOP2, RAPSN, BEST2, DPEP2 
##     HMGB2, NAPRT1, KRT8, PPIB, DSCC1, POU4F3, CCDC102A, GAPDHS, CHST6, AGXT2 
##     LYZL2, MTMR8, ACTG2, ACPL2, BANF1, PPAPDC2, HTR1A, IFI30, CYBRD1, LHX8 
## Negative:  FAIM2, CAMK2A, SCN1A, CAMK2B, FRRS1L, UNC80, PHYHIP, RASGRF2, CCK, GRIA2 
##     STXBP5L, ARPP21, SLC12A5, DIRAS2, RYR2, SLC4A10, KCNT1, GRM5, CAMKV, KIAA1211L 
##     GABRA4, GABRA1, SV2B, CX3CL1, AK5, PNMA2, JPH4, DGKG, GPR158, KCNC2 
## PC_ 2 
## Positive:  CAMKK1, DGKQ, NT5DC3, CA7, ABCG4, HTR1A, C5orf28, OTOP2, HYKK, DPEP2 
##     CHST6, POTEI, SLC8A3, SLC38A11, ADRA2A, MPPED1, MTMR8, HTR7, CACNA1B, PPAPDC2 
##     C2orf69, GRIK1, IFI30, STK32B, RASL10B, SLC24A4, FAXDC2, ADCY3, ACSS2, ANKRD29 
## Negative:  RAN, HSP90AA1, H2AFZ, HNRNPAB, CCT5, NPM1, GNG5, DBI, HMGB2, ITM2B 
##     ATP6V1G1, SERPINH1, CIRBP, CD63, NDUFA6, MDK, JUN, MYL12B, SPARC, NPC2 
##     GLUL, ID3, EEF1A1, VIM, CLIC1, COX6B1, LDHA, DDAH2, ENO1, CNN3 
## PC_ 3 
## Positive:  ADGRL2, AC011288.2, RP11-420N3.3, RP11-191L9.4, NRXN3, PLPPR1, RP11-123O10.4, ZNF385D, AC114765.1, NWD2 
##     RBFOX3, MIR137HG, MIR325HG, SGOL1-AS1, POU6F2, ANKRD18A, LY86-AS1, LINC01197, DGCR5, DPY19L1P1 
##     MIR4300HG, AQP4-AS1, HPSE2, LINC00632, NLGN4X, AC067956.1, PWRN1, LINC00599, CABP1, LINC01158 
## Negative:  KRTCAP2, APOE, C20orf24, PDIA6, PGLS, GNG11, S100A13, HIST1H2BI, ISCA2, GSTM5 
##     LAPTM4A, CST3, TMEM176B, KLF4, PDLIM2, CAP1, S100A16, APRT, CYR61, FAIM 
##     IFITM3, CDKN1A, KLF2, CLIC1, ARPC1B, IER2, S100A1, CMTM5, FXYD1, TCN2 
## PC_ 4 
## Positive:  RESP18, CTXN2, ATP6V1G2, GNG13, DISP2, C15orf59, CCDC85A, GNG3, SYNGR3, RGS8 
##     VWA5B2, C1QL3, HPCA, TUBB3, CALB1, SNCB, HTR3A, ARHGDIG, L1CAM, NAP1L5 
##     PCDH20, HMP19, DBNDD2, NPAS4, FABP3, CALY, FAM43B, CKMT1B, LOC728392, LTK 
## Negative:  PTPN18, SLCO1A2, LINC00639, INPP5D, IFI44, LYN, DISC1, NEAT1, NRGN, CMYA5 
##     IFI44L, GALNT15, PARP14, AC012593.1, AQP4-AS1, MSR1, MT2A, ISG15, SHROOM4, CABP1 
##     UACA, KCNQ1OT1, PART1, CNDP1, FAM153B, DGCR5, SOX2-OT, LINC00844, ADGRG1, LINC00599 
## PC_ 5 
## Positive:  MEST, IGFBP2, CNN3, FBXL7, NNAT, TUBB2B, GPC3, VIM, NKAIN4, ID1 
##     BMP7, CSRP2, NDN, DDAH2, GPX8, IGFBPL1, MARCKSL1, GSTM3, FBLN1, PARD3 
##     MFAP4, PTN, FABP7, COPS6, CTNNA2, ZBTB20, BEX1, CD81, ENO1, NPAS3 
## Negative:  C1QB, FCGR2A, MS4A6A, TYROBP, C1QC, AIF1, C1QA, CSF1R, CD86, MRC1 
##     MS4A7, CTSS, CCL24, FCER1G, CD53, CD14, FCGR1A, PLEK, C3AR1, LYZ 
##     FCGR2B, CX3CR1, CCL3L3, CCL2, CCR1, CD68, C5AR1, PF4, HPGDS, LY86
## No graphs detected. Computing.
## Computing nearest neighbor graph
## Computing SNN
## Using graph: RNA_snn
## + Filtering results by `var1_search`: astro
## + 6 entries matching `var1_search` identified.
## + Adding original names to results
## + Returning 50 pair-wise similarities.
cat("\nLabels similar to 'astro':\n")
## 
## Labels similar to 'astro':
print(unique(astro_neighbors$Var2))
##  [1] "fly.Aerts2020.Cortex_glia"                                  
##  [2] "fly.Aerts2020.Ensheathing_glia"                             
##  [3] "fly.Aerts2020.Perineurial_glia"                             
##  [4] "mouse.Zeisel2016.Astro2"                                    
##  [5] "mouse.Zeisel2016.Astro1"                                    
##  [6] "mouse.Zeisel2018.Subventricular.zone.radial.glia.like.cells"
##  [7] "fly.Aerts2020.X76"                                          
##  [8] "fly.Aerts2020.Plasmatocytes"                                
##  [9] "fly.Aerts2020.Subperineurial_glia"                          
## [10] "human.descartes_SampledData.Lens_fibre_cells"               
## [11] "mouse.Zeisel2016.Oligo1"                                    
## [12] "mouse.Zeisel2018.Olfactory.ensheathing.cells"               
## [13] "human.descartes_SampledData.Inhibitory_neurons"             
## [14] "human.descartes_SampledData.Amacrine_cells"                 
## [15] "human.descartes_SampledData.Horizontal_cells"               
## [16] "mouse.Zeisel2018.Dentate.gyrus.radial.glia.like.cells"      
## [17] "mouse.Zeisel2018.Ependymal.cells"                           
## [18] "human.descartes_SampledData.PDE11A_FAM19A2_positive_cells"  
## [19] "mouse.Zeisel2018.Astrocytes"                                
## [20] "mouse.DRONC_mouse.ASC1"                                     
## [21] "mouse.DRONC_mouse.ASC2"                                     
## [22] "mouse.DRONC_mouse.ChP"                                      
## [23] "mouse.Zeisel2016.Epend"                                     
## [24] "mouse.DRONC_mouse.ODC"                                      
## [25] "human.DRONC_human.NSC"                                      
## [26] "mouse.Zeisel2016.Oligo2"                                    
## [27] "mouse.Zeisel2016.Oligo6"                                    
## [28] "mouse.Zeisel2018.Oligodendrocyte.precursor.cells"           
## [29] "mouse.Zeisel2018.Oligodendrocytes"                          
## [30] "human.DRONC_human.ASC1"                                     
## [31] "human.DRONC_human.ASC2"

Strategy 3: Combined Approach

Combine TF-IDF with neighbor analysis for robust harmonization.

Identify Clusters by Enriched Terms

# Find clusters enriched for neuronal terms
neuronal_terms <- c("neuron", "neuronal", "glutamatergic", "gabaergic")

neuronal_clusters <- tfidf_df |>
    dplyr::filter(word %in% neuronal_terms) |>
    dplyr::group_by(cluster) |>
    dplyr::summarise(
        neuronal_score = sum(tf_idf),
        top_term = word[which.max(tf_idf)]
    ) |>
    dplyr::arrange(dplyr::desc(neuronal_score))
## Warning: Returning more (or less) than 1 row per `summarise()` group was deprecated in
## dplyr 1.1.0.
##  Please use `reframe()` instead.
##  When switching from `summarise()` to `reframe()`, remember that `reframe()`
##   always returns an ungrouped data frame and adjust accordingly.
## Call `lifecycle::last_lifecycle_warnings()` to see where this warning was
## generated.
## `summarise()` has grouped output by 'cluster'. You can override using the
## `.groups` argument.
print(neuronal_clusters)
## # A tibble: 3 × 3
## # Groups:   cluster [15]
##   cluster neuronal_score top_term
##   <fct>            <dbl> <chr>   
## 1 6               0.0704 neuron  
## 2 0               0.0428 neuron  
## 3 2               0.0419 neuron

Use neighbor search to validate that cells in these clusters are transcriptomically similar:

# Get cells from a neuronal cluster
if (nrow(neuronal_clusters) > 0) {
    top_cluster <- neuronal_clusters$cluster[1]
    is_top <- pseudo_seurat$cluster == top_cluster
    cluster_cells <- pseudo_seurat$celltype[is_top]
    cat("Labels in top neuronal cluster:\n")
    print(table(cluster_cells))
}
## Labels in top neuronal cluster:
## cluster_cells
##                                                 A 
##                                                 0 
##                            AFP_ALB_positive_cells 
##                                                 0 
##                                              ASC1 
##                                                 0 
##                                              ASC2 
##                                                 0 
##                                      Acinar_cells 
##                                                 0 
##                              Adrenocortical_cells 
##                                                 0 
##                                    Amacrine_cells 
##                                                 0 
##                          Antigen_presenting_cells 
##                                                 0 
##                                               Ast 
##                                                 0 
##                                              AstA 
##                                                 0 
##                                            Astro1 
##                                                 0 
##                                            Astro2 
##                                                 0 
##                                         Astrocyte 
##                                                 0 
##                                        Astrocytes 
##                                                 0 
##                                     Bipolar_cells 
##                                                 0 
##                                        Blood_e7_0 
##                                                 0 
##                                        Blood_e8_0 
##                                                 0 
##                                        Blood_e8_5 
##                                                 0 
##                                        Blood_e9_0 
##                                                 0 
##                                       Blood_e10_0 
##                                                 0 
##                                       Blood_e11_0 
##                                                 0 
##                                       Blood_e12_0 
##                                                 0 
##                                       Blood_e12_5 
##                                                 0 
##                                       Blood_e13_0 
##                                                 0 
##                                       Blood_e13_5 
##                                                 0 
##                                       Blood_e14_0 
##                                                 0 
##                                       Blood_e14_5 
##                                                 0 
##                                       Blood_e15_0 
##                                                 0 
##                                       Blood_e15_5 
##                                                 0 
##                                       Blood_e16_0 
##                                                 0 
##                                       Blood_e16_5 
##                                                 0 
##                                      Blood_e16_25 
##                                                 0 
##                                       Blood_e17_0 
##                                                 0 
##                                       Blood_e17_5 
##                                                 0 
##                                       Blood_e18_0 
##                                                 0 
##         Bronchiolar_and_alveolar_epithelial_cells 
##                                                 0 
##                                                C2 
##                                                 0 
##                                                C3 
##                                                 0 
##                                           CA1Pyr1 
##                                                 0 
##                                           CA1Pyr2 
##                                                 0 
##                                         CA1PyrInt 
##                                                 0 
##                                           CA2Pyr2 
##                                                 0 
##                                              CCAP 
##                                                 0 
##                                             CCHa1 
##                                                 0 
##                        CCL19_CCL21_positive_cells 
##                                                 0 
##                          CLC_IL5RA_positive_cells 
##                                                 0 
##                          CSH1_CSH2_positive_cells 
##                                                 0 
##                                              Capa 
##                                                 0 
##                                    Cardiomyocytes 
##                                                 0 
##                                        Cerebellum 
##                                                 1 
##                                               ChP 
##                                                 0 
##                                       Chiasm_glia 
##                                                 0 
##                                       Cholinergic 
##                                                 1 
##                                           Choroid 
##                                                 0 
##                               Choroid_plexus_e9_0 
##                                                 0 
##                              Choroid_plexus_e10_0 
##                                                 0 
##                              Choroid_plexus_e11_0 
##                                                 0 
##                              Choroid_plexus_e12_0 
##                                                 0 
##                              Choroid_plexus_e12_5 
##                                                 0 
##                              Choroid_plexus_e13_0 
##                                                 0 
##                              Choroid_plexus_e13_5 
##                                                 0 
##                              Choroid_plexus_e14_0 
##                                                 0 
##                              Choroid_plexus_e14_5 
##                                                 0 
##                              Choroid_plexus_e15_0 
##                                                 0 
##                              Choroid_plexus_e15_5 
##                                                 0 
##                              Choroid_plexus_e16_0 
##                                                 0 
##                              Choroid_plexus_e16_5 
##                                                 0 
##                             Choroid_plexus_e16_25 
##                                                 0 
##                              Choroid_plexus_e17_0 
##                                                 0 
##                              Choroid_plexus_e17_5 
##                                                 0 
##                              Choroid_plexus_e18_0 
##                                                 0 
##                                  Chromaffin_cells 
##                                                 0 
##                         Ciliated_epithelial_cells 
##                                                 0 
##                                           ClauPyr 
##                                                 0 
##                                             Clock 
##                                                 0 
##         Corneal_and_conjunctival_epithelial_cells 
##                                                 0 
##                                       Cortex_glia 
##                                                 0 
##                                               Crz 
##                                                 0 
##                                               DCN 
##                                                 0 
##                                               DN1 
##                                                 0 
##                                           Dentate 
##                                                 0 
##                                                Di 
##                                                 2 
##                                               Dm8 
##                                                 0 
##                                               Dm9 
##                                                 0 
##                                      Dopaminergic 
##                                                 0 
##                                      Ductal_cells 
##                                                 0 
##                         ELF3_AGBL2_positive_cells 
##                                                 0 
##                                               END 
##                                                 0 
##                                          ENS_glia 
##                                                 0 
##                                       ENS_neurons 
##                                                 1 
##                            Early_Neural_Tube_e7_0 
##                                                 0 
##                            Early_Neural_Tube_e8_0 
##                                                 0 
##                            Early_Neural_Tube_e8_5 
##                                                 0 
##                                     Ectoderm_e7_0 
##                                                 0 
##                                     Ectoderm_e8_0 
##                                                 0 
##                                     Ectoderm_e8_5 
##                                                 0 
##                                     Ectoderm_e9_0 
##                                                 0 
##                                    Ectoderm_e10_0 
##                                                 0 
##                                    Ectoderm_e11_0 
##                                                 0 
##                                    Ectoderm_e12_0 
##                                                 0 
##                                    Ectoderm_e12_5 
##                                                 0 
##                                    Ectoderm_e13_5 
##                                                 0 
##                                    Ectoderm_e14_5 
##                                                 0 
##                                    Ectoderm_e15_0 
##                                                 0 
##                                    Ectoderm_e15_5 
##                                                 0 
##                                               End 
##                                                 0 
##                                 Endocardial_cells 
##                                                 0 
##                                     Endoderm_e7_0 
##                                                 0 
##                                     Endoderm_e8_0 
##                                                 0 
##                                     Endoderm_e8_5 
##                                                 0 
##                                  Ensheathing_glia 
##                                                 0 
##                                           Enteric 
##                                                 1 
##                                             Epend 
##                                                 0 
##                                         Ependymal 
##                                                 0 
##                              Epicardial_fat_cells 
##                                                 0 
##                                     Erythroblasts 
##                                                 0 
##                                               Ex1 
##                                                 0 
##                                               Ex2 
##                                                 0 
##                                              Ex3a 
##                                                 0 
##                                              Ex3b 
##                                                 0 
##                                              Ex3c 
##                                                 0 
##                                              Ex3d 
##                                                 0 
##                                              Ex3e 
##                                                 0 
##                                               Ex4 
##                                                 0 
##                                              Ex5a 
##                                                 0 
##                                              Ex5b 
##                                                 0 
##                                              Ex6a 
##                                                 0 
##                                              Ex6b 
##                                                 0 
##                                               Ex8 
##                                                 0 
##                                Excitatory_neurons 
##                                                 0 
##                         Extravillous_trophoblasts 
##                                                 0 
##                                             FMRFa 
##                                                 0 
##                                   Fibroblast_e9_0 
##                                                 0 
##                                  Fibroblast_e10_0 
##                                                 0 
##                                  Fibroblast_e11_0 
##                                                 0 
##                                  Fibroblast_e12_0 
##                                                 0 
##                                  Fibroblast_e12_5 
##                                                 0 
##                                  Fibroblast_e13_0 
##                                                 0 
##                                  Fibroblast_e13_5 
##                                                 0 
##                                  Fibroblast_e14_0 
##                                                 0 
##                                  Fibroblast_e14_5 
##                                                 0 
##                                  Fibroblast_e15_0 
##                                                 0 
##                                  Fibroblast_e15_5 
##                                                 0 
##                                  Fibroblast_e16_0 
##                                                 0 
##                                  Fibroblast_e16_5 
##                                                 0 
##                                 Fibroblast_e16_25 
##                                                 0 
##                                  Fibroblast_e17_0 
##                                                 0 
##                                  Fibroblast_e17_5 
##                                                 0 
##                                  Fibroblast_e18_0 
##                                                 0 
##                                                 G 
##                                                 0 
##                                             GABA1 
##                                                 0 
##                                             GABA2 
##                                                 0 
##                                         GABAergic 
##                                                 0 
##                                    Ganglion_cells 
##                                                 0 
##                                         Glia_e9_0 
##                                                 0 
##                                        Glia_e10_0 
##                                                 0 
##                                        Glia_e11_0 
##                                                 0 
##                                        Glia_e12_0 
##                                                 0 
##                                        Glia_e12_5 
##                                                 0 
##                                        Glia_e13_0 
##                                                 0 
##                                        Glia_e13_5 
##                                                 0 
##                                        Glia_e14_0 
##                                                 0 
##                                        Glia_e14_5 
##                                                 0 
##                                        Glia_e15_0 
##                                                 0 
##                                        Glia_e15_5 
##                                                 0 
##                                        Glia_e16_0 
##                                                 0 
##                                        Glia_e16_5 
##                                                 0 
##                                       Glia_e16_25 
##                                                 0 
##                                        Glia_e17_0 
##                                                 0 
##                                        Glia_e17_5 
##                                                 0 
##                                        Glia_e18_0 
##                                                 0 
##                                     Glutamatergic 
##                                                 0 
##                                      Goblet_cells 
##                                                 0 
##                                             Gr43a 
##                                                 0 
##                                   Granule_neurons 
##                                                 0 
##                          Hematopoietic_stem_cells 
##                                                 0 
##                                      Hepatoblasts 
##                                                 0 
##                                         Hindbrain 
##                                                 1 
##                                  Horizontal_cells 
##                                                 0 
##                                               Hsp 
##                                                 0 
##                                               Hug 
##                                                 0 
##                        IGFBP1_DKK1_positive_cells 
##                                                 0 
##                                               IPC 
##                                                 0 
##                                               ITP 
##                                                 0 
##                                       Immune_e9_0 
##                                                 0 
##                                      Immune_e10_0 
##                                                 0 
##                                      Immune_e11_0 
##                                                 0 
##                                      Immune_e12_0 
##                                                 0 
##                                      Immune_e12_5 
##                                                 0 
##                                      Immune_e13_0 
##                                                 0 
##                                      Immune_e13_5 
##                                                 0 
##                                      Immune_e14_0 
##                                                 0 
##                                      Immune_e14_5 
##                                                 0 
##                                      Immune_e15_0 
##                                                 0 
##                                      Immune_e15_5 
##                                                 0 
##                                      Immune_e16_0 
##                                                 0 
##                                      Immune_e16_5 
##                                                 0 
##                                     Immune_e16_25 
##                                                 0 
##                                      Immune_e17_0 
##                                                 0 
##                                      Immune_e17_5 
##                                                 0 
##                                      Immune_e18_0 
##                                                 0 
##                                              In1a 
##                                                 0 
##                                              In1b 
##                                                 0 
##                                              In1c 
##                                                 0 
##                                               In2 
##                                                 0 
##                                               In3 
##                                                 0 
##                                              In4a 
##                                                 0 
##                                              In4b 
##                                                 0 
##                                              In6a 
##                                                 0 
##                                              In6b 
##                                                 0 
##                                               In7 
##                                                 0 
##                                               In8 
##                                                 0 
##                           Inhibitory_interneurons 
##                                                 1 
##                                Inhibitory_neurons 
##                                                 0 
##                                              Int1 
##                                                 0 
##                                              Int2 
##                                                 0 
##                                              Int3 
##                                                 0 
##                                              Int4 
##                                                 0 
##                                              Int5 
##                                                 0 
##                                              Int6 
##                                                 0 
##                                              Int7 
##                                                 0 
##                                              Int8 
##                                                 0 
##                                              Int9 
##                                                 0 
##                                             Int10 
##                                                 0 
##                                             Int11 
##                                                 0 
##                                             Int12 
##                                                 0 
##                                             Int13 
##                                                 0 
##                                             Int14 
##                                                 0 
##                                             Int15 
##                                                 0 
##                                             Int16 
##                                                 0 
##                       Intestinal_epithelial_cells 
##                                                 0 
##                             Islet_endocrine_cells 
##                                                 0 
##                                                L1 
##                                                 0 
##                                                L2 
##                                                 0 
##                                                L3 
##                                                 0 
##                                                L4 
##                                                 0 
##                                               LNv 
##                                                 0 
##                                  Lamina_monopolar 
##                                                 0 
##                                             Lawf1 
##                                                 0 
##                                             Lawf2 
##                                                 0 
##                                  Lens_fibre_cells 
##                                                 0 
##                             Limbic_system_neurons 
##                                                 0 
##                       Lymphatic_endothelial_cells 
##                                                 0 
##                                    Lymphoid_cells 
##                                                 0 
##                                              MBON 
##                                                 0 
##                                                MG 
##                                                 0 
##                        MUC13_DMBT1_positive_cells 
##                                                 0 
##                                    Megakaryocytes 
##                                                 0 
##                                   Mesangial_cells 
##                                                 0 
##                                     Mesoderm_e7_0 
##                                                 0 
##                                     Mesoderm_e8_0 
##                                                 0 
##                                     Mesoderm_e8_5 
##                                                 0 
##                                 Mesothelial_cells 
##                                                 0 
##                                 Metanephric_cells 
##                                                 0 
##                                              Mgl1 
##                                                 0 
##                                              Mgl2 
##                                                 0 
##                                               Mi1 
##                                                 0 
##                                               Mic 
##                                                 0 
##                                         Microglia 
##                                                 0 
##                                               Mip 
##                                                 0 
##                                     Myeloid_cells 
##                                                 0 
##                                               NSC 
##                                                 0 
##                                 Neural_crest_e7_0 
##                                                 0 
##                                 Neural_crest_e8_0 
##                                                 0 
##                                 Neural_crest_e8_5 
##                                                 0 
##                                 Neural_crest_e9_0 
##                                                 0 
##                                Neural_crest_e10_0 
##                                                 0 
##                                Neural_crest_e11_0 
##                                                 0 
##                              Neuroendocrine_cells 
##                                                 0 
##                                       Neuron_e8_0 
##                                                 0 
##                                       Neuron_e8_5 
##                                                 0 
##                                       Neuron_e9_0 
##                                                 0 
##                                      Neuron_e10_0 
##                                                 0 
##                                      Neuron_e11_0 
##                                                 0 
##                                      Neuron_e12_0 
##                                                 0 
##                                      Neuron_e12_5 
##                                                 1 
##                                      Neuron_e13_0 
##                                                 1 
##                                      Neuron_e13_5 
##                                                 1 
##                                      Neuron_e14_0 
##                                                 1 
##                                      Neuron_e14_5 
##                                                 0 
##                                      Neuron_e15_0 
##                                                 0 
##                                      Neuron_e15_5 
##                                                 0 
##                                      Neuron_e16_0 
##                                                 1 
##                                      Neuron_e16_5 
##                                                 0 
##                                     Neuron_e16_25 
##                                                 0 
##                                      Neuron_e17_0 
##                                                 0 
##                                      Neuron_e17_5 
##                                                 0 
##                                      Neuron_e18_0 
##                                                 0 
##                                         Node_e7_0 
##                                                 0 
##                                         Node_e8_0 
##                                                 0 
##                                               Non 
##                                                 0 
##                                               ODC 
##                                                 0 
##                                              ODC1 
##                                                 0 
##                                              ODC2 
##                                                 0 
##                                               OPC 
##                                                 0 
##                                        OPCs_e12_0 
##                                                 0 
##                                        OPCs_e12_5 
##                                                 0 
##                                        OPCs_e13_0 
##                                                 0 
##                                        OPCs_e13_5 
##                                                 0 
##                                        OPCs_e14_0 
##                                                 0 
##                                        OPCs_e14_5 
##                                                 0 
##                                        OPCs_e15_0 
##                                                 0 
##                                        OPCs_e15_5 
##                                                 0 
##                                        OPCs_e16_0 
##                                                 0 
##                                        OPCs_e16_5 
##                                                 0 
##                                       OPCs_e16_25 
##                                                 0 
##                                        OPCs_e17_0 
##                                                 0 
##                                        OPCs_e17_5 
##                                                 0 
##                                        OPCs_e18_0 
##                                                 0 
##                                    Octopaminergic 
##                                                 0 
##                                         Olfactory 
##                                                 0 
##                      Olfactory_projection_neurons 
##                                                 0 
##                                               Oli 
##                                                 0 
##                                            Oligo1 
##                                                 0 
##                                            Oligo2 
##                                                 0 
##                                            Oligo3 
##                                                 0 
##                                            Oligo4 
##                                                 0 
##                                            Oligo5 
##                                                 0 
##                                            Oligo6 
##                                                 0 
##                                   Oligo_precursor 
##                                                 0 
##                                   Oligodendrocyte 
##                                                 0 
##                                  Oligodendrocytes 
##                                                 0 
##                         PAEP_MECOM_positive_cells 
##                                                 0 
##                                               PAM 
##                                                 0 
##                        PDE1C_ACSM3_positive_cells 
##                                                 0 
##                     PDE11A_FAM19A2_positive_cells 
##                                                 0 
##                          Parietal_and_chief_cells 
##                                                 0 
##                                       Peptidergic 
##                                                 1 
##                                               Per 
##                                                 0 
##                                             Peric 
##                                                 0 
##                                         Pericytes 
##                                                 0 
##                                  Perineurial_glia 
##                                                 0 
##                                        Peripheral 
##                                                 3 
##                                      Perivascular 
##                                                 0 
##                               Photoreceptor_cells 
##                                                 0 
##                                    Photoreceptors 
##                                                 0 
##                                     Plasmatocytes 
##                                                 0 
##                       Pluripotent_stem_cells_e7_0 
##                                                 0 
##                       Pluripotent_stem_cells_e8_0 
##                                                 0 
##                       Pluripotent_stem_cells_e8_5 
##                                                 0 
##                                               Pm1 
##                                                 0 
##                                               Pm3 
##                                                 0 
##                                               Pm4 
##                                                 0 
##                                              Poxn 
##                                                 0 
##                                              Proc 
##                                                 0 
##                                  Purkinje_neurons 
##                                                 0 
##                                              Pvm1 
##                                                 0 
##                                              Pvm2 
##                                                 0 
##                                               RPE 
##                                                 0 
##                               RPE_differentiation 
##                                                 0 
##                                  Radial_glia_e9_0 
##                                                 0 
##                                 Radial_glia_e10_0 
##                                                 0 
##                                 Radial_glia_e11_0 
##                                                 0 
##                                 Radial_glia_e12_0 
##                                                 0 
##                                 Radial_glia_e12_5 
##                                                 0 
##                                 Radial_glia_e13_0 
##                                                 0 
##                                 Radial_glia_e13_5 
##                                                 0 
##                                 Radial_glia_e14_0 
##                                                 0 
##                                 Radial_glia_e14_5 
##                                                 0 
##                                 Radial_glia_e15_0 
##                                                 0 
##                                 Radial_glia_e15_5 
##                                                 0 
##                                 Radial_glia_e16_0 
##                                                 0 
##                                 Radial_glia_e16_5 
##                                                 0 
##                                Radial_glia_e16_25 
##                                                 0 
##                                 Radial_glia_e17_0 
##                                                 0 
##                                 Radial_glia_e17_5 
##                                                 0 
##                                 Radial_glia_e18_0 
##                                                 0 
##                             Retinal_pigment_cells 
##                                                 0 
##               Retinal_progenitors_and_Muller_glia 
##                                                 0 
##                                           S1PyrDL 
##                                                 0 
##                                           S1PyrL4 
##                                                 0 
##                                           S1PyrL5 
##                                                 0 
##                                          S1PyrL5a 
##                                                 0 
##                                           S1PyrL6 
##                                                 0 
##                                          S1PyrL6b 
##                                                 0 
##                                          S1PyrL23 
##                                                 0 
##                        SATB2_LRRC7_positive_cells 
##                                                 0 
##                        SKOR2_NPSR1_positive_cells 
##                                                 0 
##                      SLC24A4_PEX5L_positive_cells 
##                                                 0 
##                       SLC26A4_PAEP_positive_cells 
##                                                 0 
##                                               SMC 
##                                                 0 
##                          STC2_TLX1_positive_cells 
##                                                 0 
##                                         Satellite 
##                                                 0 
##                                   Satellite_cells 
##                                                 0 
##                                     Schwann_cells 
##                                                 0 
##                                     Schwann_e12_0 
##                                                 0 
##                                     Schwann_e13_0 
##                                                 0 
##                                     Schwann_e13_5 
##                                                 0 
##                                     Schwann_e14_0 
##                                                 0 
##                                     Schwann_e14_5 
##                                                 0 
##                                     Schwann_e15_0 
##                                                 0 
##                                     Schwann_e15_5 
##                                                 0 
##                                     Schwann_e16_0 
##                                                 0 
##                                     Schwann_e16_5 
##                                                 0 
##                                    Schwann_e16_25 
##                                                 0 
##                                     Schwann_e17_0 
##                                                 0 
##                                     Schwann_e17_5 
##                                                 0 
##                                     Schwann_e18_0 
##                                                 0 
##                                      Serotonergic 
##                                                 0 
##                             Skeletal_muscle_cells 
##                                                 0 
##                               Smooth_muscle_cells 
##                                                 0 
##                                            Spinal 
##                                                 2 
##                         Squamous_epithelial_cells 
##                                                 0 
##                                    Stellate_cells 
##                                                 0 
##                                     Stromal_cells 
##                                                 0 
##                                            SubPyr 
##                                                 0 
##                               Subperineurial_glia 
##                                                 0 
##                                    Subventricular 
##                                                 0 
##                                       Sympathetic 
##                                                 2 
##                                    Sympathoblasts 
##                                                 0 
## Syncytiotrophoblasts_and_villous_cytotrophoblasts 
##                                                 0 
##                                                T1 
##                                                 0 
##                                                T2 
##                                                 0 
##                                                T3 
##                                                 0 
##                                                T4 
##                                                 0 
##                                     Telencephalon 
##                                                 0 
##                           Thymic_epithelial_cells 
##                                                 0 
##                                        Thymocytes 
##                                                 0 
##                                               Tm1 
##                                                 0 
##                                             Tm5ab 
##                                                 0 
##                                              Tm5c 
##                                                 0 
##                                               Tm9 
##                                                 0 
##                                             TmY14 
##                                                 0 
##                                Trophectoderm_e7_0 
##                                                 0 
##                                Trophectoderm_e8_0 
##                                                 0 
##                                Trophectoderm_e8_5 
##                                                 0 
##                           Trophoblast_giant_cells 
##                                                 0 
##                                      Tyraminergic 
##                                                 0 
##                              Unipolar_brush_cells 
##                                                 0 
##                                Ureteric_bud_cells 
##                                                 0 
##                                          Vascular 
##                                                 0 
##                                     Vascular_e7_0 
##                                                 0 
##                                     Vascular_e8_0 
##                                                 0 
##                                     Vascular_e8_5 
##                                                 0 
##                                     Vascular_e9_0 
##                                                 0 
##                                    Vascular_e10_0 
##                                                 0 
##                                    Vascular_e11_0 
##                                                 0 
##                                    Vascular_e12_0 
##                                                 0 
##                                    Vascular_e12_5 
##                                                 0 
##                                    Vascular_e13_0 
##                                                 0 
##                                    Vascular_e13_5 
##                                                 0 
##                                    Vascular_e14_0 
##                                                 0 
##                                    Vascular_e14_5 
##                                                 0 
##                                    Vascular_e15_0 
##                                                 0 
##                                    Vascular_e15_5 
##                                                 0 
##                                    Vascular_e16_0 
##                                                 0 
##                                    Vascular_e16_5 
##                                                 0 
##                                   Vascular_e16_25 
##                                                 0 
##                                    Vascular_e17_0 
##                                                 0 
##                                    Vascular_e17_5 
##                                                 0 
##                                    Vascular_e18_0 
##                                                 0 
##                        Vascular_endothelial_cells 
##                                                 0 
##                                             Vend1 
##                                                 0 
##                                             Vend2 
##                                                 0 
##                                  Visceral_neurons 
##                                                 1 
##                                              Vsmc 
##                                                 0 
##                                                 X 
##                                                 0 
##                                                X0 
##                                                 0 
##                                                X1 
##                                                 0 
##                                                X2 
##                                                 0 
##                                                X3 
##                                                 0 
##                                                X4 
##                                                 0 
##                                                X5 
##                                                 0 
##                                                X6 
##                                                 0 
##                                                X7 
##                                                 0 
##                                                X9 
##                                                 0 
##                                               X13 
##                                                 0 
##                                               X15 
##                                                 0 
##                                               X16 
##                                                 0 
##                                               X18 
##                                                 0 
##                                               X19 
##                                                 0 
##                                               X21 
##                                                 0 
##                                               X25 
##                                                 0 
##                                               X29 
##                                                 0 
##                                               X30 
##                                                 0 
##                                               X32 
##                                                 0 
##                                               X34 
##                                                 0 
##                                               X40 
##                                                 0 
##                                               X44 
##                                                 0 
##                                               X47 
##                                                 0 
##                                               X49 
##                                                 0 
##                                               X50 
##                                                 0 
##                                               X51 
##                                                 0 
##                                               X54 
##                                                 0 
##                                               X56 
##                                                 0 
##                                               X59 
##                                                 0 
##                                               X62 
##                                                 0 
##                                               X67 
##                                                 0 
##                                               X68 
##                                                 0 
##                                               X69 
##                                                 0 
##                                               X71 
##                                                 0 
##                                               X75 
##                                                 0 
##                                               X76 
##                                                 0 
##                                               X78 
##                                                 0 
##                                               X79 
##                                                 0 
##                                               X81 
##                                                 0 
##                                               X83 
##                                                 0 
##                                               X84 
##                                                 0 
##                                              adPN 
##                                                 0 
##                                      adaxial_cell 
##                                                 0 
##                                  blood_precursors 
##                                                 0 
##                                         cartilage 
##                                                 0 
##                                        cerebellum 
##                                                 0 
##                              comitted_progenitors 
##                                                 0 
##                             comittted_progenitors 
##                                                 0 
##                             committed_progenitors 
##                                                 0 
##                                            cornea 
##                                                 0 
##                                  cranial_ganglion 
##                                                 2 
##                                     cycling_cells 
##                                                 0 
##                               cycling_progenitors 
##                                                 0 
##                                       dermal_bone 
##                                                 0 
##                                             dienc 
##                                                 0 
##                                      diencephalon 
##                                                 1 
##                                        dorsal_Fan 
##                                                 0 
##                               dorsal_diencephalon 
##                                                 0 
##                            dorsal_diencephalon_II 
##                                                 0 
##                              dorsal_dienchephalon 
##                                                 0 
##                                   dorsal_habenula 
##                                                 2 
##                              dorsal_telencephalon 
##                                                 2 
##                                epidermal_ionocyte 
##                                                 0 
##                                         epidermis 
##                                                 0 
##                                        epithelium 
##                                                 0 
##                                      erythrocytes 
##                                                 0 
##                                             exCA1 
##                                                 0 
##                                             exCA3 
##                                                 0 
##                                              exDG 
##                                                 0 
##                                            exPFC1 
##                                                 0 
##                                            exPFC2 
##                                                 0 
##                                            exPFC3 
##                                                 0 
##                                            exPFC4 
##                                                 0 
##                                            exPFC5 
##                                                 0 
##                                            exPFC6 
##                                                 0 
##                                            exPFC7 
##                                                 0 
##                                            exPFC8 
##                                                 0 
##                                            exPFC9 
##                                                 0 
##                                               eye 
##                                                 0 
##                                       floor_plate 
##                                                 0 
##                                              glia 
##                                                 0 
##                                  glia_progenitors 
##                                                 0 
##                                 glial_progenitors 
##                                                 0 
##                                     granule_cells 
##                                                 1 
##                                  heart_primordium 
##                                                 0 
##                             hinbrain_rhombomere_5 
##                                                 0 
##                             hinbrain_rhombomere_6 
##                                                 0 
##                             hinbrain_rhombomere_7 
##                                                 0 
##                                         hindbrain 
##                                                 0 
##                            hindbrain_rhombomere_1 
##                                                 0 
##                            hindbrain_rhombomere_2 
##                                                 0 
##                            hindbrain_rhombomere_3 
##                                                 0 
##                            hindbrain_rhombomere_4 
##                                                 0 
##                            hindbrain_rhombomere_5 
##                                                 0 
##                            hindbrain_rhombomere_6 
##                                                 0 
##                            hindbrain_rhombomere_7 
##                                                 0 
##                                      hypothalamus 
##                                                 1 
##                                               lPN 
##                                                 0 
##                                              lens 
##                                                 0 
##                              lens_differentiation 
##                                                 0 
##                                   lens_epithelium 
##                                                 0 
##                                       macrophages 
##                                                 0 
##                                          mesoderm 
##                                                 0 
##                                         microglia 
##                                                 0 
##                                               mid 
##                                                 1 
##                                          midbrain 
##                                                 1 
##                               midbrain_neural_rod 
##                                                 0 
##                                            muscle 
##                                                 0 
##                                      neural_crest 
##                                                 0 
##                              neural_crest_derived 
##                                                 0 
##                                         neuromast 
##                                                 0 
##                                           neurons 
##                                                12 
##                                       neutrophils 
##                                                 0 
##                                         notochord 
##                                                 0 
##                                    olfactory_bulb 
##                                                 0 
##                                  oligodendrocytes 
##                                                 0 
##                                       optic_stalk 
##                                                 0 
##                                      optic_tectum 
##                                                 1 
##                                     optic_vesicle 
##                                                 0 
##                                              otic 
##                                                 0 
##                                      otic_vesicle 
##                                                 0 
##                                      pectoral_fin 
##                                                 0 
##                                pectoral_fin_field 
##                                                 0 
##                                          periderm 
##                                                 0 
##                                   pharangeal_arch 
##                                                 0 
##                                    pharyneal_arch 
##                                                 0 
##                                   pharyngeal_arch 
##                                                 0 
##                               pharyngeal_endoderm 
##                                                 0 
##                    pharyngeal_erythrocyte_lineage 
##                                                 0 
##                                      pigment_cell 
##                                                 0 
##                                           placode 
##                                                 0 
##                                     possibly_otic 
##                                                 0 
##                                  prechordal_plate 
##                                                 0 
##                                       progenitors 
##                                                 1 
##                                        pronephros 
##                                                 0 
##                                  purkinje_neurons 
##                                                 2 
##                                       radial_glia 
##                                                 0 
##                                            retina 
##                                                 2 
##                                retina_neuroblasts 
##                                                 0 
##                               retina_neurogenesis 
##                                                 0 
##                                    retina_neurons 
##                                                 1 
##                              rostral_blood_island 
##                                                 0 
##                                   sensory_neurons 
##                                                 0 
##                                            somite 
##                                                 0 
##                                     telencephalon 
##                                                 2 
##                                   telencephalon_I 
##                                                 0 
##                                  telencephalon_II 
##                                                 0 
##                              ventral_diencephalon 
##                                                 0 
##                                 ventral_forebrain 
##                                                 0 
##                                  ventral_habenula 
##                                                 0 
##                                  ventral_midbrain 
##                                                 0

Visualizing Cross-dataset Agreement

Color by Dataset

Visualize how cells from different datasets intermix:

res_batch <- plot_tfidf(
    obj = pseudo_seurat,
    label_var = "celltype",
    cluster_var = "cluster",
    color_var = "batch",
    show_plot = TRUE
)
## Extracting obsm from Seurat: umap
## + Dropping 2 conflicting obs variables: UMAP.1, UMAP.2
## Setting cell metadata (obs) in obj.
## Warning in ggplot2::geom_point(ggplot2::aes_string(color = color_var, size =
## size_var, : Ignoring unknown aesthetics: label

Clusters with cells from multiple datasets that share enriched terms indicate successful cross-dataset harmonization.

Color by Species

For cross-species comparisons:

res_species <- plot_tfidf(
    obj = pseudo_seurat,
    label_var = "celltype",
    cluster_var = "cluster",
    color_var = "species",
    show_plot = TRUE
)
## Extracting obsm from Seurat: umap
## + Dropping 2 conflicting obs variables: UMAP.1, UMAP.2
## Setting cell metadata (obs) in obj.
## Warning in ggplot2::geom_point(ggplot2::aes_string(color = color_var, size =
## size_var, : Ignoring unknown aesthetics: label

Word Cloud Summary

Generate a word cloud to summarize the most distinctive terms across all clusters:

wc <- wordcloud_tfidf(
    obj = pseudo_seurat,
    label_var = "celltype",
    cluster_var = "cluster",
    terms_per_cluster = 5
)
## Loading required namespace: ggwordcloud
## Extracting obsm from Seurat: umap
## + Dropping 2 conflicting obs variables: UMAP.1, UMAP.2
## Setting cell metadata (obs) in obj.
## Warning in ggplot2::geom_point(ggplot2::aes_string(color = color_var, size =
## size_var, : Ignoring unknown aesthetics: label

Practical Recommendations

  1. Start with TF-IDF: Identify enriched terms to understand the semantic content of each cluster.

  2. Use neighbor search for validation: Confirm that cells with similar labels are transcriptomically similar.

  3. Examine cross-dataset mixing: Well-harmonized cell types should have cells from multiple datasets in the same cluster.

  4. Iterate: Refine cluster assignments and re-run TF-IDF to improve harmonization.

  5. Document mappings: Create a mapping table of original labels to harmonized labels for reproducibility.

Session Info

utils::sessionInfo()
## R Under development (unstable) (2026-01-22 r89323)
## Platform: x86_64-pc-linux-gnu
## Running under: Ubuntu 24.04.3 LTS
## 
## Matrix products: default
## BLAS:   /usr/lib/x86_64-linux-gnu/openblas-pthread/libblas.so.3 
## LAPACK: /usr/lib/x86_64-linux-gnu/openblas-pthread/libopenblasp-r0.3.26.so;  LAPACK version 3.12.0
## 
## locale:
##  [1] LC_CTYPE=en_US.UTF-8       LC_NUMERIC=C              
##  [3] LC_TIME=en_US.UTF-8        LC_COLLATE=en_US.UTF-8    
##  [5] LC_MONETARY=en_US.UTF-8    LC_MESSAGES=en_US.UTF-8   
##  [7] LC_PAPER=en_US.UTF-8       LC_NAME=C                 
##  [9] LC_ADDRESS=C               LC_TELEPHONE=C            
## [11] LC_MEASUREMENT=en_US.UTF-8 LC_IDENTIFICATION=C       
## 
## time zone: UTC
## tzcode source: system (glibc)
## 
## attached base packages:
## [1] stats     graphics  grDevices utils     datasets  methods   base     
## 
## other attached packages:
## [1] future_1.69.0    scNLP_0.99.0     BiocStyle_2.39.0
## 
## loaded via a namespace (and not attached):
##   [1] RColorBrewer_1.1-3     jsonlite_2.0.0         magrittr_2.0.4        
##   [4] spatstat.utils_3.2-1   farver_2.1.2           rmarkdown_2.30        
##   [7] fs_1.6.6               ragg_1.5.0             vctrs_0.7.1           
##  [10] ROCR_1.0-12            spatstat.explore_3.7-0 htmltools_0.5.9       
##  [13] janeaustenr_1.0.0      sass_0.4.10            sctransform_0.4.3     
##  [16] parallelly_1.46.1      KernSmooth_2.23-26     bslib_0.9.0           
##  [19] htmlwidgets_1.6.4      tokenizers_0.3.0       desc_1.4.3            
##  [22] ica_1.0-3              plyr_1.8.9             plotly_4.12.0         
##  [25] zoo_1.8-15             cachem_1.1.0           commonmark_2.0.0      
##  [28] igraph_2.2.1           mime_0.13              lifecycle_1.0.5       
##  [31] pkgconfig_2.0.3        Matrix_1.7-4           R6_2.6.1              
##  [34] fastmap_1.2.0          fitdistrplus_1.2-6     shiny_1.12.1          
##  [37] digest_0.6.39          tidytext_0.4.3         colorspace_2.1-2      
##  [40] patchwork_1.3.2        Seurat_5.4.0           tensor_1.5.1          
##  [43] RSpectra_0.16-2        irlba_2.3.5.1          SnowballC_0.7.1       
##  [46] textshaping_1.0.4      labeling_0.4.3         progressr_0.18.0      
##  [49] spatstat.sparse_3.1-0  httr_1.4.7             polyclip_1.10-7       
##  [52] abind_1.4-8            compiler_4.6.0         withr_3.0.2           
##  [55] S7_0.2.1               fastDummies_1.7.5      maps_3.4.3            
##  [58] MASS_7.3-65            tools_4.6.0            lmtest_0.9-40         
##  [61] otel_0.2.0             httpuv_1.6.16          future.apply_1.20.1   
##  [64] goftest_1.2-3          glue_1.8.0             nlme_3.1-168          
##  [67] promises_1.5.0         gridtext_0.1.5         grid_4.6.0            
##  [70] Rtsne_0.17             cluster_2.1.8.1        reshape2_1.4.5        
##  [73] generics_0.1.4         isoband_0.3.0          gtable_0.3.6          
##  [76] spatstat.data_3.1-9    tidyr_1.3.2            data.table_1.18.0     
##  [79] xml2_1.5.2             utf8_1.2.6             sp_2.2-0              
##  [82] spatstat.geom_3.7-0    RcppAnnoy_0.0.23       markdown_2.0          
##  [85] ggrepel_0.9.6          RANN_2.6.2             pillar_1.11.1         
##  [88] stringr_1.6.0          pals_1.10              spam_2.11-3           
##  [91] RcppHNSW_0.6.0         later_1.4.5            splines_4.6.0         
##  [94] dplyr_1.1.4            lattice_0.22-7         survival_3.8-6        
##  [97] deldir_2.0-4           tidyselect_1.2.1       miniUI_0.1.2          
## [100] pbapply_1.7-4          knitr_1.51             gridExtra_2.3         
## [103] litedown_0.9           bookdown_0.46          scattermore_1.2       
## [106] xfun_0.56              matrixStats_1.5.0      stringi_1.8.7         
## [109] lazyeval_0.2.2         yaml_2.3.12            evaluate_1.0.5        
## [112] codetools_0.2-20       ggwordcloud_0.6.2      tibble_3.3.1          
## [115] BiocManager_1.30.27    cli_3.6.5              uwot_0.2.4            
## [118] xtable_1.8-4           reticulate_1.44.1      systemfonts_1.3.1     
## [121] jquerylib_0.1.4        dichromat_2.0-0.1      Rcpp_1.1.1            
## [124] globals_0.18.0         spatstat.random_3.4-4  mapproj_1.2.12        
## [127] png_0.1-8              spatstat.univar_3.1-6  parallel_4.6.0        
## [130] pkgdown_2.2.0          ggplot2_4.0.1          dotCall64_1.2         
## [133] listenv_0.10.0         viridisLite_0.4.2      scales_1.4.0          
## [136] ggridges_0.5.7         SeuratObject_5.3.0     purrr_1.2.1           
## [139] rlang_1.1.7            cowplot_1.2.0