Detect features that are spatially correlated. Functions for starting from either a gobject (detectSpatialCorFeats()) or individual pieces of data (detectSpatialCorFeatsMatrix()) are provided.

detectSpatialCorFeats(
  gobject,
  spat_unit = NULL,
  feat_type = NULL,
  spat_loc_name = "raw",
  method = c("grid", "network"),
  expression_values = c("normalized", "scaled", "custom"),
  subset_feats = NULL,
  spatial_network_name = "Delaunay_network",
  network_smoothing = NULL,
  spatial_grid_name = "spatial_grid",
  min_cells_per_grid = 4,
  cor_method = c("pearson", "kendall", "spearman")
)

detectSpatialCorFeatsMatrix(
  expression_matrix,
  method = c("grid", "network"),
  spatial_network,
  spatial_grid,
  spatial_locs,
  subset_feats = NULL,
  network_smoothing = NULL,
  min_cells_per_grid = 4,
  cor_method = c("pearson", "kendall", "spearman")
)

Arguments

gobject

giotto object

spat_unit

spatial unit

feat_type

feature type

spat_loc_name

name for spatial locations

method

method to use for spatial averaging

expression_values

gene expression values to use

subset_feats

subset of features to use

spatial_network_name

name of spatial network to use

network_smoothing

smoothing factor between 0 and 1 (has automatic default, see details)

spatial_grid_name

name of spatial grid to use

min_cells_per_grid

minimum number of cells to consider a grid

cor_method

correlation method

expression_matrix

provided expression matrix

spatial_network

provided spatial network

spatial_grid

provided spatial grid

spatial_locs

provided spatial locations

Value

returns a spatial correlation object: spatCorObject

Details

For method = network, it expects a fully connected spatial network. You can make sure to create a fully connected network by setting minimal_k > 0 in the createSpatialNetwork function.

  1. grid-averaging: average gene expression values within a predefined spatial grid

  2. network-averaging: smoothens the gene expression matrix by averaging the expression within one cell by using the neighbours within the predefined spatial network. \(b\) is a smoothening factor passed by network_smoothing param that defaults to \(1 - 1/k\), where \(k\) is the median number of k-neighbors in the selected spatial network. Setting \(b = 0\) means no smoothing and \(b = 1\) means no contribution from its own expression.

The spatCorObject can be further explored with showSpatialCorFeats()

Examples

g <- GiottoData::loadGiottoMini("visium")
#> 1. read Giotto object
#> 2. read Giotto feature information
#> 3. read Giotto spatial information
#> 3.1 read Giotto spatial shape information
#> 3.2 read Giotto spatial centroid information
#> 3.3 read Giotto spatial overlap information
#> 4. read Giotto image information
#> python already initialized in this session
#>  active environment : '/usr/bin/python3'
#>  python version : 3.10
#> checking default envname 'giotto_env'
#> a system default python environment was found
#> Using python path:
#>  "/usr/bin/python3"

# Perform with data in a gobject
detectSpatialCorFeats(g, method = "network")
#> $cor_DT
#>               feat_ID      variable   spat_cor   expr_cor     cordiff spatrank
#>                <char>        <fctr>      <num>      <num>       <num>    <int>
#>      1: 1110017D15Rik 1110017D15Rik  1.0000000  1.0000000  0.00000000        1
#>      2: 1110017D15Rik        Dnali1  0.5152451  0.3690438  0.14620131        2
#>      3: 1110017D15Rik         Spag8  0.5044653  0.3180572  0.18640810        3
#>      4: 1110017D15Rik 1700001C02Rik  0.4957575  0.4021766  0.09358091        4
#>      5: 1110017D15Rik       Ccdc153  0.4947645  0.4050736  0.08969090        5
#>     ---                                                                       
#> 401952:         Zmat4         Cabp7 -0.3106553 -0.2099353 -0.10072002      630
#> 401953:         Zmat4           Sst -0.3310337 -0.1903481 -0.14068568      631
#> 401954:         Zmat4          Hopx -0.3886083 -0.2587739 -0.12983435      632
#> 401955:         Zmat4           Ttr -0.4280274 -0.3482754 -0.07975206      633
#> 401956:         Zmat4          Gfap -0.4757224 -0.3562773 -0.11944508      634
#>         exprrank rankdiff
#>            <int>    <int>
#>      1:        1        0
#>      2:        6       -4
#>      3:       10       -7
#>      4:        3        1
#>      5:        2        3
#>     ---                  
#> 401952:      630        0
#> 401953:      627        4
#> 401954:      632        0
#> 401955:      633        0
#> 401956:      634        0
#> 
#> $feat_order
#>   [1] "Gna12"         "Ccnd2"         "Btbd17"        "Sox9"         
#>   [5] "Sez6"          "Serpinf1"      "S100a6"        "Col1a1"       
#>   [9] "Vwf"           "Esam"          "Npas1"         "Tiam1"        
#>  [13] "Bcam"          "Hmgn2"         "Rab3b"         "Inmt"         
#>  [17] "Ddr1"          "Homer3"        "Cp"            "Calb2"        
#>  [21] "Man1a"         "Efnb3"         "Hlf"           "Gstm7"        
#>  [25] "Cavin1"        "Sst"           "Hapln2"        "Hspb1"        
#>  [29] "Ndrg1"         "Nid1"          "Kit"           "Pvalb"        
#>  [33] "Tmbim1"        "Crip1"         "Tek"           "Elovl1"       
#>  [37] "Cyba"          "Itih3"         "Cnp"           "Hap1"         
#>  [41] "Clic1"         "Stx1a"         "Arpp19"        "Dio2"         
#>  [45] "Id3"           "Crlf1"         "Ttc9b"         "Carhsp1"      
#>  [49] "Cabp7"         "Fam163b"       "Rarres2"       "Syn2"         
#>  [53] "Ick"           "Prox1"         "Adssl1"        "Vipr2"        
#>  [57] "Gltp"          "Amotl1"        "Bcas1"         "Aldh1a2"      
#>  [61] "Tppp3"         "Sirt2"         "Lpl"           "Cers2"        
#>  [65] "Qdpr"          "Stk32c"        "Lbp"           "Hsd11b1"      
#>  [69] "Vtn"           "Stac2"         "Plxdc1"        "Igfbp4"       
#>  [73] "Dbndd2"        "Pltp"          "Slc12a4"       "Cadps2"       
#>  [77] "Erbb3"         "Pmp22"         "Kcnab3"        "Myh11"        
#>  [81] "Plod1"         "Etnppl"        "Arhgef25"      "Rcn3"         
#>  [85] "Vip"           "Utrn"          "Grm1"          "Reep3"        
#>  [89] "Fabp7"         "Dcn"           "Slc17a8"       "Dusp6"        
#>  [93] "Kitl"          "Ccn2"          "Timp3"         "Ascl1"        
#>  [97] "Unc5b"         "Vsir"          "Gamt"          "Ptprb"        
#> [101] "Csrp2"         "Adarb1"        "Rnf130"        "Nefh"         
#> [105] "Aebp1"         "Sept4"         "Gria1"         "Sypl"         
#> [109] "Arsg"          "Cacng4"        "Rab37"         "Itgb4"        
#> [113] "Aspa"          "Cygb"          "Rflnb"         "Doc2b"        
#> [117] "Gfap"          "Npas3"         "Serpina3n"     "Clmn"         
#> [121] "Prkch"         "Fbln5"         "Akr1c18"       "Gng4"         
#> [125] "Sema4d"        "Cxcl14"        "Hexb"          "Iqgap2"       
#> [129] "Crhbp"         "Rab3c"         "Thbs4"         "Serinc5"      
#> [133] "Erbin"         "Prkcd"         "Pdlim2"        "Ednrb"        
#> [137] "Cpne6"         "Sema5a"        "Matn2"         "Nptxr"        
#> [141] "Enpp2"         "Efcab6"        "Ppp1r1a"       "Litaf"        
#> [145] "Il1rap"        "Apod"          "Cpox"          "Grik1"        
#> [149] "Clic6"         "Nr4a1"         "Igfbp6"        "Sncg"         
#> [153] "Gzma"          "Grm2"          "Prph"          "Rsph1"        
#> [157] "Sox8"          "Fam234a"       "Lims2"         "Aqp4"         
#> [161] "Slc12a2"       "Cd74"          "Pdgfrb"        "Csf1r"        
#> [165] "Cbln2"         "Frmd8"         "Gldc"          "Tcf7l2"       
#> [169] "Bhlhe22"       "Kcnip2"        "Alas2"         "Gdf11"        
#> [173] "Baiap2"        "Ifitm3"        "Phkg1"         "Nptx1"        
#> [177] "Pde8a"         "Itih5"         "Homer2"        "Fam20c"       
#> [181] "Cplx2"         "Myo5b"         "Snca"          "Col3a1"       
#> [185] "Ecrg4"         "Map4k4"        "2010300C02Rik" "Ogfrl1"       
#> [189] "Fn1"           "Efhd1"         "Ngef"          "Bok"          
#> [193] "Tmem163"       "Lct"           "Cfh"           "Ptpn4"        
#> [197] "Tnni1"         "Csrp1"         "Nfasc"         "Syt2"         
#> [201] "Atp2b4"        "Rgs16"         "Ncf2"          "Cnih3"        
#> [205] "Tmem63a"       "Cfap126"       "Meig1"         "Ccdc3"        
#> [209] "Rgs5"          "Enkur"         "Myoc"          "Vim"          
#> [213] "Prkcq"         "Gad2"          "Lcn2"          "Nr4a2"        
#> [217] "Ermn"          "Fibcd1"        "Gsn"           "Grb14"        
#> [221] "Notch1"        "Ube2l6"        "Gatm"          "Cd82"         
#> [225] "Mdk"           "Lamp5"         "Itpka"         "Chgb"         
#> [229] "Mal"           "Pdyn"          "Nkx2-2"        "Edn3"         
#> [233] "Car2"          "Col9a3"        "Chrna4"        "Gss"          
#> [237] "Slc7a11"       "Tm4sf1"        "Kcnab1"        "Shox2"        
#> [241] "Serpini1"      "Tspan2"        "S100a11"       "Vcam1"        
#> [245] "Lef1"          "Npy2r"         "F3"            "Calb1"        
#> [249] "Epha7"         "Slc44a1"       "1110017D15Rik" "Tpm2"         
#> [253] "Slc6a9"        "Laptm5"        "Sema3c"        "Hpca"         
#> [257] "Padi2"         "Slc4a2"        "Gabrd"         "Rgs12"        
#> [261] "Slc30a3"       "Emilin1"       "1700001C02Rik" "Gabra4"       
#> [265] "Pdgfra"        "Epha5"         "Tesc"          "Pitpnm2"      
#> [269] "Mmp17"         "Cit"           "Foxp2"         "Arpc1b"       
#> [273] "Col1a2"        "Eln"           "Pcolce"        "Tsc22d4"      
#> [277] "Cald1"         "Ccdc136"       "Npy"           "Slc13a4"      
#> [281] "Gkn3"          "Slc6a13"       "Mgp"           "Lmo3"         
#> [285] "Slco1c1"       "Cpne9"         "Itpr2"         "Slc6a11"      
#> [289] "Vamp1"         "Cd9"           "Pglyrp1"       "Slc17a6"      
#> [293] "Sipa1l3"       "Sh3gl3"        "Plekhb1"       "Nupr1"        
#> [297] "Dkk3"          "Cox6a2"        "Dkkl1"         "Fgfr2"        
#> [301] "Crym"          "Slc38a5"       "Flna"          "Bgn"          
#> [305] "Col4a1"        "Plat"          "Nr3c2"         "Cbln1"        
#> [309] "Gab1"          "Pllp"          "Necab2"        "Dbndd1"       
#> [313] "Agt"           "Cryab"         "Tagln"         "Mcam"         
#> [317] "Olfm2"         "Icam5"         "Anxa2"         "Rora"         
#> [321] "Calml4"        "Htr3a"         "Paqr5"         "Rasgrf1"      
#> [325] "Zic1"          "Mobp"          "Hhatl"         "Trf"          
#> [329] "Cpne4"         "Cdhr4"         "Cd59a"         "Gng11"        
#> [333] "Chrm1"         "Prr5l"         "Ugt8a"         "Rims3"        
#> [337] "Sgpp2"         "Camkv"         "Sox10"         "Micall1"      
#> [341] "Tie1"          "Fa2h"          "Ucp2"          "Vav3"         
#> [345] "Ccp110"        "Gria2"         "Rasl10a"       "Foxj1"        
#> [349] "Trim59"        "Dynlrb2"       "Gjc1"          "Neurod1"      
#> [353] "Ttyh2"         "Cpne7"         "Nrip3"         "Tbr1"         
#> [357] "Lcat"          "Tmem98"        "Myo1d"         "Prdm8"        
#> [361] "Tgfbi"         "Kcnc2"         "Thrsp"         "Isg15"        
#> [365] "Acta2"         "H2-Q4"         "Dock4"         "Myrf"         
#> [369] "Lmo1"          "Igfbp7"        "P2ry12"        "Pcdh8"        
#> [373] "Lgi4"          "Fxyd1"         "Fxyd7"         "Mag"          
#> [377] "Zcchc12"       "Anln"          "Wnt4"          "C1ql2"        
#> [381] "Unc93b1"       "Tspan15"       "Ppp1r14a"      "Clic4"        
#> [385] "Galnt6"        "Ccn3"          "Rims2"         "Zmat4"        
#> [389] "Cldn11"        "Ccdc33"        "Slc32a1"       "Tgm2"         
#> [393] "Vstm2l"        "Neurod6"       "Crygn"         "Enpp6"        
#> [397] "Neurod2"       "Satb2"         "Pcp4l1"        "Txnip"        
#> [401] "Egr1"          "Otud7b"        "Car14"         "Rgs4"         
#> [405] "Dock10"        "Lpar1"         "Trps1"         "Ptpn3"        
#> [409] "Lefty1"        "Zfhx3"         "St6galnac5"    "Ak5"          
#> [413] "Hrh3"          "Nexn"          "Adgrl4"        "Rlbp1"        
#> [417] "Igfbp2"        "Wfs1"          "Cplx3"         "Orai2"        
#> [421] "Olig2"         "Gpr37"         "Plcb4"         "Nxph4"        
#> [425] "Ndufa4l2"      "Cacng5"        "Plekhg1"       "Cmtm5"        
#> [429] "Gpr161"        "Rfx3"          "Igsf21"        "Ramp3"        
#> [433] "Arhgap12"      "Nkx6-2"        "Ninj2"         "Cldn5"        
#> [437] "Htr2c"         "Hspb8"         "Fmod"          "Prelp"        
#> [441] "Mpped1"        "Rapgef5"       "Vwa1"          "Stab1"        
#> [445] "Arl15"         "Reln"          "Mustn1"        "Pbxip1"       
#> [449] "Dnali1"        "Id1"           "Lrrtm3"        "Tmem212"      
#> [453] "Epop"          "Gjc2"          "1190005I06Rik" "Ctla2a"       
#> [457] "Cnr1"          "Gpr4"          "Sowaha"        "Tent5c"       
#> [461] "Serpinb1a"     "S1pr5"         "Penk"          "Cdc42ep2"     
#> [465] "Slitrk6"       "Hpcal4"        "Olig1"         "Rprml"        
#> [469] "Stxbp6"        "Hs3st2"        "Rbp1"          "Ppm1e"        
#> [473] "Sox18"         "Kctd4"         "Nxph3"         "Vat1l"        
#> [477] "Cd24a"         "Odf3b"         "Gjb1"          "Selplg"       
#> [481] "Bcl11b"        "Lhfp"          "Ranbp3l"       "Msx1"         
#> [485] "Zdhhc22"       "Phldb1"        "Ctxn1"         "Tprn"         
#> [489] "Vstm2a"        "Ndnf"          "Pdp1"          "Scn3b"        
#> [493] "Cdc42ep1"      "Gal3st1"       "Rasd1"         "Spink8"       
#> [497] "Opalin"        "Gja4"          "Synpo2"        "Pcdh20"       
#> [501] "Adra1b"        "Fam131a"       "Tmem125"       "Mb21d2"       
#> [505] "Lingo3"        "Ncald"         "Siglech"       "Rgs14"        
#> [509] "Gm14964"       "Gpr17"         "Cx3cr1"        "Ezr"          
#> [513] "Adarb2"        "Cpne8"         "Plekhg3"       "Lamb2"        
#> [517] "Creb5"         "Cntn2"         "Aldh1a1"       "Phgdh"        
#> [521] "Tnfaip6"       "Camk2d"        "Shisa6"        "Cthrc1"       
#> [525] "Tmem119"       "Hmgb2"         "Tmem158"       "Dsp"          
#> [529] "Lrtm2"         "Gabra5"        "Tns1"          "Nell1"        
#> [533] "Slain1"        "Prr18"         "S100a8"        "S100a9"       
#> [537] "Col8a2"        "Synpr"         "Sertm1"        "Adgrf5"       
#> [541] "Capg"          "Gjc3"          "Fcer1g"        "Pde1a"        
#> [545] "Tafa1"         "Ddn"           "Hopx"          "Ptk2b"        
#> [549] "Rgs3"          "Kcng2"         "Ntng1"         "H2-Q7"        
#> [553] "Ifitm2"        "Plaat3"        "Plekhh1"       "Myl4"         
#> [557] "H2-K1"         "Dpp6"          "Tmem91"        "Cyp2d22"      
#> [561] "Ttr"           "Suclg2"        "Patj"          "Btbd3"        
#> [565] "Unc13c"        "Syt9"          "Shisal1"       "Grm4"         
#> [569] "Sox11"         "Slc22a8"       "Slc24a3"       "Klk8"         
#> [573] "Ipcef1"        "Tnnt1"         "Hhip"          "Spag8"        
#> [577] "Fxyd6"         "Ppp1r3c"       "Cbln4"         "Nnat"         
#> [581] "Myl9"          "Vxn"           "Lgals1"        "Gpr88"        
#> [585] "Ctxn3"         "Lyz2"          "Tmem100"       "Wfdc17"       
#> [589] "Ahnak"         "Hba-a2"        "Hba-a1"        "Ccdc153"      
#> [593] "Ccnd1"         "Serpinh1"      "Gad1"          "Arhgef10"     
#> [597] "2410004P03Rik" "Nt5dc2"        "Cebpd"         "Sod3"         
#> [601] "H2-D1"         "C4b"           "Tmem88b"       "Hbb-bt"       
#> [605] "Rhog"          "Rprm"          "Ly6a"          "Mog"          
#> [609] "Trbc2"         "Igkc"          "Smim1"         "Hs3st4"       
#> [613] "Evi2a"         "Prkcg"         "Ly6c1"         "Gbp4"         
#> [617] "Wipf3"         "BC039966"      "Gm13889"       "Plcxd2"       
#> [621] "Mia"           "Pou3f1"        "Lrrc10b"       "Apol11b"      
#> [625] "Gpr62"         "Shisa8"        "Gm2115"        "Kctd12"       
#> [629] "1700047M11Rik" "Lhfpl3"        "C030029H02Rik" "Gm19935"      
#> [633] "9630013A20Rik" "2900040C04Rik"
#> 
#> $cor_hclust
#> list()
#> 
#> $cor_clusters
#> list()
#> 
#> attr(,"class")
#> [1] "spatCorObject" "list"         

# This analysis can also be performed with data outside of the gobject
detectSpatialCorFeatsMatrix(
    expression_matrix = getExpression(
        g,
        output = "matrix"
    ),
    method = "network",
    spatial_network = getSpatialNetwork(g, output = "networkDT")
)
#> $cor_DT
#>               feat_ID      variable   spat_cor    expr_cor     cordiff spatrank
#>                <char>        <fctr>      <num>       <num>       <num>    <int>
#>      1: 1110017D15Rik 1110017D15Rik  1.0000000  1.00000000  0.00000000        1
#>      2: 1110017D15Rik       Dynlrb2  0.9071450  0.84973343  0.05741162        2
#>      3: 1110017D15Rik       Ccdc153  0.9054963  0.84504575  0.06045057        3
#>      4: 1110017D15Rik       Tmem212  0.9024137  0.81451232  0.08790135        4
#>      5: 1110017D15Rik       Cfap126  0.8814460  0.77851057  0.10293547        5
#>     ---                                                                        
#> 401952:         Zmat4         Thbs4 -0.1808436 -0.08644981 -0.09439376      630
#> 401953:         Zmat4           Mal -0.1814648 -0.10522798 -0.07623678      631
#> 401954:         Zmat4          Fa2h -0.1874828 -0.08735847 -0.10012430      632
#> 401955:         Zmat4          Gfap -0.1943739 -0.13487620 -0.05949770      633
#> 401956:         Zmat4         Ninj2 -0.2061581 -0.11224072 -0.09391739      634
#>         exprrank rankdiff
#>            <int>    <int>
#>      1:        1        0
#>      2:        2        0
#>      3:        3        0
#>      4:        4        0
#>      5:        8       -3
#>     ---                  
#> 401952:      620       10
#> 401953:      632       -1
#> 401954:      622       10
#> 401955:      634       -1
#> 401956:      633        1
#> 
#> $feat_order
#>   [1] "Gna12"         "Ccnd2"         "Btbd17"        "Sox9"         
#>   [5] "Sez6"          "Serpinf1"      "S100a6"        "Col1a1"       
#>   [9] "Vwf"           "Esam"          "Npas1"         "Tiam1"        
#>  [13] "Bcam"          "Hmgn2"         "Rab3b"         "Inmt"         
#>  [17] "Ddr1"          "Homer3"        "Cp"            "Calb2"        
#>  [21] "Man1a"         "Efnb3"         "Hlf"           "Gstm7"        
#>  [25] "Cavin1"        "Sst"           "Hapln2"        "Hspb1"        
#>  [29] "Ndrg1"         "Nid1"          "Kit"           "Pvalb"        
#>  [33] "Tmbim1"        "Crip1"         "Tek"           "Elovl1"       
#>  [37] "Cyba"          "Itih3"         "Cnp"           "Hap1"         
#>  [41] "Clic1"         "Stx1a"         "Arpp19"        "Dio2"         
#>  [45] "Id3"           "Crlf1"         "Ttc9b"         "Carhsp1"      
#>  [49] "Cabp7"         "Fam163b"       "Rarres2"       "Syn2"         
#>  [53] "Ick"           "Prox1"         "Adssl1"        "Vipr2"        
#>  [57] "Gltp"          "Amotl1"        "Bcas1"         "Aldh1a2"      
#>  [61] "Tppp3"         "Sirt2"         "Lpl"           "Cers2"        
#>  [65] "Qdpr"          "Stk32c"        "Lbp"           "Hsd11b1"      
#>  [69] "Vtn"           "Stac2"         "Plxdc1"        "Igfbp4"       
#>  [73] "Dbndd2"        "Pltp"          "Slc12a4"       "Cadps2"       
#>  [77] "Erbb3"         "Pmp22"         "Kcnab3"        "Myh11"        
#>  [81] "Plod1"         "Etnppl"        "Arhgef25"      "Rcn3"         
#>  [85] "Vip"           "Utrn"          "Grm1"          "Reep3"        
#>  [89] "Fabp7"         "Dcn"           "Slc17a8"       "Dusp6"        
#>  [93] "Kitl"          "Ccn2"          "Timp3"         "Ascl1"        
#>  [97] "Unc5b"         "Vsir"          "Gamt"          "Ptprb"        
#> [101] "Csrp2"         "Adarb1"        "Rnf130"        "Nefh"         
#> [105] "Aebp1"         "Sept4"         "Gria1"         "Sypl"         
#> [109] "Arsg"          "Cacng4"        "Rab37"         "Itgb4"        
#> [113] "Aspa"          "Cygb"          "Rflnb"         "Doc2b"        
#> [117] "Gfap"          "Npas3"         "Serpina3n"     "Clmn"         
#> [121] "Prkch"         "Fbln5"         "Akr1c18"       "Gng4"         
#> [125] "Sema4d"        "Cxcl14"        "Hexb"          "Iqgap2"       
#> [129] "Crhbp"         "Rab3c"         "Thbs4"         "Serinc5"      
#> [133] "Erbin"         "Prkcd"         "Pdlim2"        "Ednrb"        
#> [137] "Cpne6"         "Sema5a"        "Matn2"         "Nptxr"        
#> [141] "Enpp2"         "Efcab6"        "Ppp1r1a"       "Litaf"        
#> [145] "Il1rap"        "Apod"          "Cpox"          "Grik1"        
#> [149] "Clic6"         "Nr4a1"         "Igfbp6"        "Sncg"         
#> [153] "Gzma"          "Grm2"          "Prph"          "Rsph1"        
#> [157] "Sox8"          "Fam234a"       "Lims2"         "Aqp4"         
#> [161] "Slc12a2"       "Cd74"          "Pdgfrb"        "Csf1r"        
#> [165] "Cbln2"         "Frmd8"         "Gldc"          "Tcf7l2"       
#> [169] "Bhlhe22"       "Kcnip2"        "Alas2"         "Gdf11"        
#> [173] "Baiap2"        "Ifitm3"        "Phkg1"         "Nptx1"        
#> [177] "Pde8a"         "Itih5"         "Homer2"        "Fam20c"       
#> [181] "Cplx2"         "Myo5b"         "Snca"          "Col3a1"       
#> [185] "Ecrg4"         "Map4k4"        "2010300C02Rik" "Ogfrl1"       
#> [189] "Fn1"           "Efhd1"         "Ngef"          "Bok"          
#> [193] "Tmem163"       "Lct"           "Cfh"           "Ptpn4"        
#> [197] "Tnni1"         "Csrp1"         "Nfasc"         "Syt2"         
#> [201] "Atp2b4"        "Rgs16"         "Ncf2"          "Cnih3"        
#> [205] "Tmem63a"       "Cfap126"       "Meig1"         "Ccdc3"        
#> [209] "Rgs5"          "Enkur"         "Myoc"          "Vim"          
#> [213] "Prkcq"         "Gad2"          "Lcn2"          "Nr4a2"        
#> [217] "Ermn"          "Fibcd1"        "Gsn"           "Grb14"        
#> [221] "Notch1"        "Ube2l6"        "Gatm"          "Cd82"         
#> [225] "Mdk"           "Lamp5"         "Itpka"         "Chgb"         
#> [229] "Mal"           "Pdyn"          "Nkx2-2"        "Edn3"         
#> [233] "Car2"          "Col9a3"        "Chrna4"        "Gss"          
#> [237] "Slc7a11"       "Tm4sf1"        "Kcnab1"        "Shox2"        
#> [241] "Serpini1"      "Tspan2"        "S100a11"       "Vcam1"        
#> [245] "Lef1"          "Npy2r"         "F3"            "Calb1"        
#> [249] "Epha7"         "Slc44a1"       "1110017D15Rik" "Tpm2"         
#> [253] "Slc6a9"        "Laptm5"        "Sema3c"        "Hpca"         
#> [257] "Padi2"         "Slc4a2"        "Gabrd"         "Rgs12"        
#> [261] "Slc30a3"       "Emilin1"       "1700001C02Rik" "Gabra4"       
#> [265] "Pdgfra"        "Epha5"         "Tesc"          "Pitpnm2"      
#> [269] "Mmp17"         "Cit"           "Foxp2"         "Arpc1b"       
#> [273] "Col1a2"        "Eln"           "Pcolce"        "Tsc22d4"      
#> [277] "Cald1"         "Ccdc136"       "Npy"           "Slc13a4"      
#> [281] "Gkn3"          "Slc6a13"       "Mgp"           "Lmo3"         
#> [285] "Slco1c1"       "Cpne9"         "Itpr2"         "Slc6a11"      
#> [289] "Vamp1"         "Cd9"           "Pglyrp1"       "Slc17a6"      
#> [293] "Sipa1l3"       "Sh3gl3"        "Plekhb1"       "Nupr1"        
#> [297] "Dkk3"          "Cox6a2"        "Dkkl1"         "Fgfr2"        
#> [301] "Crym"          "Slc38a5"       "Flna"          "Bgn"          
#> [305] "Col4a1"        "Plat"          "Nr3c2"         "Cbln1"        
#> [309] "Gab1"          "Pllp"          "Necab2"        "Dbndd1"       
#> [313] "Agt"           "Cryab"         "Tagln"         "Mcam"         
#> [317] "Olfm2"         "Icam5"         "Anxa2"         "Rora"         
#> [321] "Calml4"        "Htr3a"         "Paqr5"         "Rasgrf1"      
#> [325] "Zic1"          "Mobp"          "Hhatl"         "Trf"          
#> [329] "Cpne4"         "Cdhr4"         "Cd59a"         "Gng11"        
#> [333] "Chrm1"         "Prr5l"         "Ugt8a"         "Rims3"        
#> [337] "Sgpp2"         "Camkv"         "Sox10"         "Micall1"      
#> [341] "Tie1"          "Fa2h"          "Ucp2"          "Vav3"         
#> [345] "Ccp110"        "Gria2"         "Rasl10a"       "Foxj1"        
#> [349] "Trim59"        "Dynlrb2"       "Gjc1"          "Neurod1"      
#> [353] "Ttyh2"         "Cpne7"         "Nrip3"         "Tbr1"         
#> [357] "Lcat"          "Tmem98"        "Myo1d"         "Prdm8"        
#> [361] "Tgfbi"         "Kcnc2"         "Thrsp"         "Isg15"        
#> [365] "Acta2"         "H2-Q4"         "Dock4"         "Myrf"         
#> [369] "Lmo1"          "Igfbp7"        "P2ry12"        "Pcdh8"        
#> [373] "Lgi4"          "Fxyd1"         "Fxyd7"         "Mag"          
#> [377] "Zcchc12"       "Anln"          "Wnt4"          "C1ql2"        
#> [381] "Unc93b1"       "Tspan15"       "Ppp1r14a"      "Clic4"        
#> [385] "Galnt6"        "Ccn3"          "Rims2"         "Zmat4"        
#> [389] "Cldn11"        "Ccdc33"        "Slc32a1"       "Tgm2"         
#> [393] "Vstm2l"        "Neurod6"       "Crygn"         "Enpp6"        
#> [397] "Neurod2"       "Satb2"         "Pcp4l1"        "Txnip"        
#> [401] "Egr1"          "Otud7b"        "Car14"         "Rgs4"         
#> [405] "Dock10"        "Lpar1"         "Trps1"         "Ptpn3"        
#> [409] "Lefty1"        "Zfhx3"         "St6galnac5"    "Ak5"          
#> [413] "Hrh3"          "Nexn"          "Adgrl4"        "Rlbp1"        
#> [417] "Igfbp2"        "Wfs1"          "Cplx3"         "Orai2"        
#> [421] "Olig2"         "Gpr37"         "Plcb4"         "Nxph4"        
#> [425] "Ndufa4l2"      "Cacng5"        "Plekhg1"       "Cmtm5"        
#> [429] "Gpr161"        "Rfx3"          "Igsf21"        "Ramp3"        
#> [433] "Arhgap12"      "Nkx6-2"        "Ninj2"         "Cldn5"        
#> [437] "Htr2c"         "Hspb8"         "Fmod"          "Prelp"        
#> [441] "Mpped1"        "Rapgef5"       "Vwa1"          "Stab1"        
#> [445] "Arl15"         "Reln"          "Mustn1"        "Pbxip1"       
#> [449] "Dnali1"        "Id1"           "Lrrtm3"        "Tmem212"      
#> [453] "Epop"          "Gjc2"          "1190005I06Rik" "Ctla2a"       
#> [457] "Cnr1"          "Gpr4"          "Sowaha"        "Tent5c"       
#> [461] "Serpinb1a"     "S1pr5"         "Penk"          "Cdc42ep2"     
#> [465] "Slitrk6"       "Hpcal4"        "Olig1"         "Rprml"        
#> [469] "Stxbp6"        "Hs3st2"        "Rbp1"          "Ppm1e"        
#> [473] "Sox18"         "Kctd4"         "Nxph3"         "Vat1l"        
#> [477] "Cd24a"         "Odf3b"         "Gjb1"          "Selplg"       
#> [481] "Bcl11b"        "Lhfp"          "Ranbp3l"       "Msx1"         
#> [485] "Zdhhc22"       "Phldb1"        "Ctxn1"         "Tprn"         
#> [489] "Vstm2a"        "Ndnf"          "Pdp1"          "Scn3b"        
#> [493] "Cdc42ep1"      "Gal3st1"       "Rasd1"         "Spink8"       
#> [497] "Opalin"        "Gja4"          "Synpo2"        "Pcdh20"       
#> [501] "Adra1b"        "Fam131a"       "Tmem125"       "Mb21d2"       
#> [505] "Lingo3"        "Ncald"         "Siglech"       "Rgs14"        
#> [509] "Gm14964"       "Gpr17"         "Cx3cr1"        "Ezr"          
#> [513] "Adarb2"        "Cpne8"         "Plekhg3"       "Lamb2"        
#> [517] "Creb5"         "Cntn2"         "Aldh1a1"       "Phgdh"        
#> [521] "Tnfaip6"       "Camk2d"        "Shisa6"        "Cthrc1"       
#> [525] "Tmem119"       "Hmgb2"         "Tmem158"       "Dsp"          
#> [529] "Lrtm2"         "Gabra5"        "Tns1"          "Nell1"        
#> [533] "Slain1"        "Prr18"         "S100a8"        "S100a9"       
#> [537] "Col8a2"        "Synpr"         "Sertm1"        "Adgrf5"       
#> [541] "Capg"          "Gjc3"          "Fcer1g"        "Pde1a"        
#> [545] "Tafa1"         "Ddn"           "Hopx"          "Ptk2b"        
#> [549] "Rgs3"          "Kcng2"         "Ntng1"         "H2-Q7"        
#> [553] "Ifitm2"        "Plaat3"        "Plekhh1"       "Myl4"         
#> [557] "H2-K1"         "Dpp6"          "Tmem91"        "Cyp2d22"      
#> [561] "Ttr"           "Suclg2"        "Patj"          "Btbd3"        
#> [565] "Unc13c"        "Syt9"          "Shisal1"       "Grm4"         
#> [569] "Sox11"         "Slc22a8"       "Slc24a3"       "Klk8"         
#> [573] "Ipcef1"        "Tnnt1"         "Hhip"          "Spag8"        
#> [577] "Fxyd6"         "Ppp1r3c"       "Cbln4"         "Nnat"         
#> [581] "Myl9"          "Vxn"           "Lgals1"        "Gpr88"        
#> [585] "Ctxn3"         "Lyz2"          "Tmem100"       "Wfdc17"       
#> [589] "Ahnak"         "Hba-a2"        "Hba-a1"        "Ccdc153"      
#> [593] "Ccnd1"         "Serpinh1"      "Gad1"          "Arhgef10"     
#> [597] "2410004P03Rik" "Nt5dc2"        "Cebpd"         "Sod3"         
#> [601] "H2-D1"         "C4b"           "Tmem88b"       "Hbb-bt"       
#> [605] "Rhog"          "Rprm"          "Ly6a"          "Mog"          
#> [609] "Trbc2"         "Igkc"          "Smim1"         "Hs3st4"       
#> [613] "Evi2a"         "Prkcg"         "Ly6c1"         "Gbp4"         
#> [617] "Wipf3"         "BC039966"      "Gm13889"       "Plcxd2"       
#> [621] "Mia"           "Pou3f1"        "Lrrc10b"       "Apol11b"      
#> [625] "Gpr62"         "Shisa8"        "Gm2115"        "Kctd12"       
#> [629] "1700047M11Rik" "Lhfpl3"        "C030029H02Rik" "Gm19935"      
#> [633] "9630013A20Rik" "2900040C04Rik"
#> 
#> $cor_hclust
#> list()
#> 
#> $cor_clusters
#> list()
#> 
#> attr(,"class")
#> [1] "list"          "spatCorObject"