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. 2020 Jun 10;40(24):4622–4643. doi: 10.1523/JNEUROSCI.2339-19.2020

Table 4.

Microglial density and microglia/neuron ratio in different cortical sitesa

Gray matter (microglia/mg)
White matter (microglia/mg)
Wilcoxon I/mg versus white matter
I/mg
M vs. I, (exponent and p value) I/N
I/mg
M vs. I, (exponent and p value)
(range and mean ± SE) (I/mg vs I, Spearman ρ and p value) (I/mg vs N/mg, Spearman ρ andp value) (I/mg vs N/mg, exponent and p value) (I/mg vs N/mg, exponent, and p value) (I vs I/mg, exponent, and p value) (range and mean ± SE) (I/mg vs I, Spearman ρ, andp value) (I/mg vs O/mg, Spearman ρ, andp value) (I/mg vs O/mg exponent, and p value)
All clades [106-16 061]
3620 ± 63
ρ=0.2942
p < 0.0001
ρ=−0.1108
p = 0.0005
−0.025 ± 0.016
p = 0.1165
0.882 ± 0.010
p < 0.0001
−1.020 ± 0.016
p < 0.0001
1.104 ± 0.063
p < 0.0001
[271-19 827]
5016 ± 93
ρ=0.1614
p < 0.0001
ρ=0.2172
p < 0.0001
0.596 ± 0.064
p < 0.0001
0.945 ± 0.012
p < 0.0001
p < 0.0001
Per clade
Afrotheria [930-7971]
4140 ± 97
ρ=0.3005
p = 0.0009
ρ=0.1538
p = 0.0936
0.083 ± 0.066
p = 0.2094
0.760 ± 0.050
p < 0.0001
−0.917 ± 0.066
p < 0.0001
0.840 ± 0.126
p < 0.0001
[371-10 673]
4482 ± 163
ρ=0.4252
p < 0.0001
ρ=0.5710
p < 0.0001
1.144 ± 0.149
p < 0.0001
0.628 ± 0.054
p < 0.0001
p = 0.2835
Artiodactyla [1075-15 647]
4975 ± 345
ρ=0.4382
p = 0.0006
ρ=0.5049
p = 0.0001
0.715 ± 0.136
p < 0.0001
0.847 ± 0.071
p < 0.0001
−0.285 ± 0.136
p = 0.0397
0.536 ± 0.088
p < 0.0001
[2576-16 518]
6833 ± 387
ρ=0.2862
p = 0.0309
ρ=0.7154
p < 0.0001
1.472 ± 0.202
p < 0.0001
0.882 ± 0.039
p < 0.0001
p = 0.0002
Carnivora [919-10 728]
3722 ± 98
ρ=0.2386
p = 0.0003
ρ=0.2954
p < 0.0001
0.122 ± 0.026
p < 0.0001
0.863 ± 0.025
p < 0.0001
−0.878 ± 0.026
p < 0.0001
0.276 ± 0.156
p = 0.0775
[741-19 827]
6740 ± 242
ρ=0.3512
p < 0.0001
ρ=0.7023
p < 0.0001
1.323 ± 0.103
p < 0.0001
0.844 ± 0.028
p < 0.0001
p < 0.0001
Marsupialia [600-15 098]
4648 ± 175
ρ=0.2954
p = 0.0003
ρ=0.3813
p < 0.0001
0.368 ± 0.092
p = 0.0001
0.791 ± 0.040
p < 0.0001
−0.632 ± 0.092
p < 0.0001
0.730 ± 0.068
p < 0.0001
[1220-12 132]
5242 ± 162
ρ=0.2362
p = 0.0080
ρ=0.4062
p < 0.0001
0.474 ± 0.128
p = 0.0003
0.890 ± 0.026
p < 0.0001
p = 0.0151
Primata [106-16 061]
2890 ± 92
ρ=0.1876
p = 0.0001
ρ=0.3226
p < 0.0001
0.301 ± 0.036
p < 0.0001
0.893 ± 0.022
p < 0.0001
−0.657 ± 0.037
p < 0.0001
0.537 ± 0.054
p < 0.0001
[271-12 511]
3965 ± 123
ρ=−0.0689
p = 0.2197
ρ=0.3998
p < 0.0001
0.864 ± 0.088
p < 0.0001
1.019 ± 0.027
p < 0.0001
p < 0.0001
Per species
Afrotheria*
L. africana [930-7971]
4170 ± 97
ρ=0.2704
p = 0.0032
ρ=0.1538
p = 0.0936
0.280 ± 0.080
p = 0.0006
0.791 ± 0.054
p < 0.0001
−0.719 ± 0.080
p < 0.0001
0.652 ± 0.099
p < 0.0001
[371-10 673]
4511 ± 166
ρ=0.4081
p < 0.0001
ρ=0.5491
p = 0.0001
1.240 ± 0.154
p < 0.0001
0.604 ± 0.058
p < 0.0001
p = 0.2732
Artiodactyla
G. camelopardalis [2764-9975]
5825 ± 352
ρ=0.6361
p = 0.0026
ρ=0.2060
p = 0.3835
0.228 ± 0.170
p = 0.1969
0.604 ± 0.141
p = 0.0004
−0.772 ± 0.170
p = 0.0003
0.602 ± 0.297
p = 0.0576
[3769-10 746]
7197 ± 466
ρ=0.5789
p = 0.0075
ρ=0.4226
p = 0.0634
0.827 ± 0.365
p = 0.0361
0.876 ± 0.101
p < 0.0001
p = 0.0223
T. strepsiceros [1075-4298]
2847 ± 214
ρ=0.6118
p = 0.0118
ρ=0.6824
p = 0.0036
1.066 ± 0.211
p = 0.0002
0.658 ± 0.133
p = 0.0002
0.066 ± 0.211
p = 0.7581
0.394 ± 0.120
p = 0.0054
[3502-6804]
5070 ± 231
ρ=0.5353
p = 0.0326
ρ=0.4471
p = 0.0825
0.432 ± 0.373
p = 0.2663
0.933 ± 0.032
p < 0.0001
p < 0.0001
S. scrofa domesticus [5584-15 647]
8481 ± 939
ρ=0.1879
p = 0.6032
ρ=−0.1758
p = 0.6272
0.400 ± 0.250
p = 0.1478
1.170 ± 0.089
p < 0.0001
−0.600 ± 0.250
p = 0.0430
0.393 ± 0.379
p = 0.3305
[8595-16 518]
11 742 ± 803
ρ=0.5333
p = 0.1392
ρ=0.5000
p = 0.1705
0.983 ± 0.525
p = 0.1032
0.907 ± 0.139
p = 0.0003
p = 0.0101
A. marsupialis [1941-6161]
3476 ± 377
ρ=−0.1189
p = 0.7129
ρ=0.8531
p = 0.0004
0.625 ± 0.114
p = 0.0003
1.138 ± 0.164
p < 0.0001
−0.375 ± 0.114
p = 0.0080
−0.203 ± 0.218
p = 0.3754
[2576-7398]
4894 ± 399
ρ=0.4685
p = 0.1245
ρ=0.9231
p < 0.0001
1.285 ± 0.220
p = 0.0002
0.831 ± 0.041
p < 0.0001
p = 0.0194
Carnivora
M. mongo [919-2130]
1400 ± 172
ρ=0.3214
p = 0.4821
ρ=0.7857
p = 0.0362
0.698 ± 0.201
p = 0.0177
0.398 ± 0.511
p = 0.4715
−0.302 ± 0.201
p = 0.1926
−0.014 ± 0.292
p = 0.9642
[740-2918]
1926 ± 156
ρ=0.7343
p = 0.0065
ρ=0.1678
p = 0.6021
1.391 ± 0.916
p = 0.1600
0.250 ± 0.193
p = 0.2235
p = 0.0572
U. arctos [1432-5774]
2801 ± 94
ρ=0.1660
p = 0.1828
ρ=−0.0423
p = 0.7359
0.022 ± 0.088
p = 0.8050
0.953 ± 0.032
p < 0.0001
−0.978 ± 0.088
p < 0.0001
0.956 ± 0.178
p < 0.0001
[1970-11 965]
5738 ± 281
ρ=0.3883
p = 0.0058
ρ=0.5573
p < 0.0001
1.178 ± 0.184
p < 0.0001
0.953 ± 0.047
p < 0.0001
p < 0.0001
F. catus [1536-4755]
2902 ± 186
ρ=0.3510
p = 0.0726
ρ=0.4829
p = 0.0107
0.588 ± 0.211
p = 0.0101
0.846 ± 0.078
p < 0.0001
−0.412 ± 0.211
p = 0.0622
0.598 ± 0.145
p = 0.0004
[2276-5590]
3406 ± 227
ρ=0.6941
p = 0.0029
ρ=0.3824
p = 0.1439
0.492 ± 0.376
p = 0.2117
0.850 ± 0.070
p < 0.0001
p = 0.0899
C. familiaris [1867-6866]
4170 ± 132
ρ=0.2634
p = 0.0646
ρ=0.1200
p = 0.4065
0.085 ± 0.076
p = 0.2666
0.948 ± 0.041
p < 0.0001
−0.915 ± 0.076
p < 0.0001
0.699 ± 0.268
p = 0.0121
[4846-15 717]
8094 ± 358
ρ=0.2406
p = 0.1773
ρ=0.6086
p = 0.0002
0.668 ± 0.142
p < 0.0001
1.018 ± 0.048
p < 0.0001
p < 0.0001
M. putorius furo [4523-10 728]
7366 ± 525
ρ=0.1636
p = 0.6307
ρ=0.4455
p = 0.1697
0.260 ± 0.160
p = 0.1396
0.892 ± 0.190
p = 0.0011
−0.740 ± 0.160
p = 0.0013
0.131 ± 0.536
p = 0.8119
[10 348-19 827]
13 278 ± 909
ρ=−0.0833
p = 0.8312
ρ=−0.1333
p = 0.7324
−0.583 ± 0.562
p = 0.3343
1.039 ± 0.144
p = 0.0002
p = 0.0003
P. leo [2192-8166]
4285 ± 126
ρ=0.2756
p = 0.0263
ρ=0.0137
p = 0.9139
0.081 ± 0.070
p = 0.2482
0.895 ± 0.051
p < 0.0001
−0.919 ± 0.070
p < 0.0001
0.740 ± 0.223
p = 0.0015
[5816-10 588]
8123 ± 199
ρ=0.4866
p = 0.0013
ρ=0.3747
p = 0.0158
0.472 ± 0.132
p = 0.0010
0.943 ± 0.026
p < 0.0001
p < 0.0001
Marsupialia
M. rufogriseus [4090-7414]
5753 ± 225
ρ=0.0610
p = 0.7927
ρ=0.1117
p = 0.6298
0.058 ± 0.075
p = 0.4500
1.053 ± 0.087
p < 0.0001
−0.942 ± 0.075
p < 0.0001
0.476 ± 0.679
p = 0.4919
[5303-9640]
6775 ± 257
ρ=−0.0561
p = 0.8194
ρ=0.2614
p = 0.2797
0.224 ± 0.247
p = 0.3770
1.053 ± 0.053
p < 0.0001
p = 0.0127
D. aurita [3290-8565]
5801 ± 425
ρ=0.8576
p < 0.0001
ρ=0.6182
p = 0.0063
0.485 ± 0.173
p = 0.0126
0.461 ± 0.100
p = 0.0003
−0.515 ± 0.173
p = 0.0088
0.319 ± 0.242
p = 0.2069
[4113-8611]
5506 ± 402
ρ=0.1678
p = 0.6021
ρ=0.6573
p = 0.0202
0.553 ± 0.131
p = 0.0018
0.884 ± 0.166
p = 0.0003
p = 0.5677
D. goodfellowi [1780-7000]
3465 ± 239
ρ=0.2591
p = 0.2214
ρ=0.2417
p = 0.2551
0.275 ± 0.168
p = 0.1174
0.991 ± 0.076
p < 0.0001
−0.725 ± 0.168
p = 0.0003
0.608 ± 0.241
p = 0.0192
[3536-7873]
5382 ± 246
ρ=0.4442
p = 0.0437
ρ=0.1558
p = 0.4999
0.259 ± 0.272
p = 0.3529
0.826 ± 0.068
p < 0.0001
p < 0.0001
M. parma [2351-15 098]
5547 ± 640
ρ=−0.0912
p = 0.7103
ρ=0.7123
p = 0.0006
0.510 ± 0.164
p = 0.0063
1.117 ± 0.174
p < 0.0001
−0.490 ± 0.164
p = 0.0082
0.287 ± 0.228
p = 0.2257
[2671-5518]
4316 ± 222
ρ=0.4071
p = 0.1320
ρ=0.5179
p = 0.0480
0.484 ± 0.248
p = 0.0730
0.831 ± 0.067
p < 0.0001
p = 0.1270
W. bicolor [1163-8680]
6191 ± 330
ρ=0.3468
p = 0.1049
ρ=−0.0524
p = 0.8124
−0.136 ± 0.209
p = 0.5223
0.736 ± 0.076
p < 0.0001
−1.136 ± 0.209
p < 0.0001
1.145 ± 0.223
p = 0.000044
[4194-12 132]
6966 ± 429
ρ=0.2403
p = 0.2942
ρ=0.5026
p = 0.0202
0.238 ± 0.179
p = 0.1987
0.937 ± 0.066
p < 0.0001
p = 0.6052
S. harrisii [600-2571]
1394 ± 156
ρ=0.3187
p = 0.2886
ρ=0.5495
p = 0.0518
0.945 ± 0.411
p = 0.0422
0.807 ± 0.226
p = 0.0045
−0.055 ± 0.411
p = 0.8956
0.657 ± 0.149
p = 0.0011
[1220-4633]
2971 ± 324
ρ=0.7273
p = 0.0074
ρ=0.3007
p = 0.3423
0.483 ± 0.634
p = 0.4638
0.765 ± 0.146
p = 0.0004
p = 0.0006
M. fuliginosus [2113-5441]
3725 ± 197
ρ=0.5047
p = 0.0062
ρ=0.2906
p = 0.1335
0.328 ± 0.171
p = 0.0657
0.916 ± 0.060
p < 0.0001
−0.672 ± 0.171
p = 0.0006
0.621 ± 0.197
p = 0.0041
[1945-5161]
4029 ± 166
ρ=0.2538
p = 0.2208
ρ=0.5646
p = 0.0033
0.881 ± 0.301
p = 0.0076
0.832 ± 0.026
p < 0.0001
p = 0.1277
Primata
P. anubiscynocephalus [856-8907]
2402 ± 154
ρ=0.0651
p = 0.6337
ρ=0.2333
p = 0.0865
0.200 ± 0.087
p = 0.0254
1.023 ± 0.050
p < 0.0001
−0.800 ± 0.087
p < 0.0001
0.546 ± 0.197
p = 0.0077
[1983-11 992]
4385 ± 320
ρ=0.2047
p = 0.1724
ρ=0.6150
p < 0.0001
1.071 ± 0.169
p < 0.0001
0.960 ± 0.103
p < 0.0001
p < 0.0001
M. fascicularis [989-11 057]
3718 ± 257
ρ=0.6443
p < 0.0001
ρ=0.4089
p = 0.0097
0.482 ± 0.178
p = 0.0102
0.756 ± 0.048
p < 0.0001
−0.518 ± 0.178
p = 0.0061
0.657 ± 0.127
p < 0.0001
[1620-10 255]
5997 ± 348
ρ=0.3559
p = 0.0306
ρ=0.1566
p = 0.3764
0.095 ± 0.220
p = 0.6700
0.873 ± 0.059
p < 0.0001
p < 0.0001
O. garnettii [3384-16 061]
9259 ± 909
ρ=0.2264
p = 0.4364
ρ=0.0637
p = 0.8286
0.021 ± 0.291
p = 0.9434
0.472 ± 0.265
p = 0.1001
−0.979 ± 0.291
p = 0.0057
0.979 ± 0.286
p = 0.0050
[2226-12 240]
7912 ± 1172
ρ=0.8833
p = 0.0016
ρ=0.5000
p = 0.1705
1.251 ± 0.341
p = 0.0080
−0.063 ± 0.221
p = 0.7835
p = 0.3951
H. sapiens sapiens [664-7812]
2379 ± 81
ρ=0.3915
p < 0.0001
ρ=−0.1368
p = 0.0695
−0.138 ± 0.058
p = 0.0192
0.889 ± 0.023
p < 0.0001
−0.993 ± 0.065
p < 0.0001
1.222 ± 0.087
p < 0.0001
[915-9079]
2891 ± 133
ρ=0.1703
p = 0.0713
ρ=0.6150
p < 0.0001
1.244 ± 0.201
p < 0.0001
0.999 ± 0.050
p < 0.0001
p = 0.0005
A. trivirgatus [1155-4145]
2612 ± 180
ρ=0.4946
p = 0.0120
ρ=0.2215
p = 0.2872
0.099 ± 0.121
p = 0.4215
0.815 ± 0.090
p < 0.0001
−0.901 ± 0.121
p < 0.0001
0.714 ± 0.350
p = 0.0530
[1782-4366]
2863 ± 193
ρ=0.1909
p = 0.4479
ρ=0.5872
p = 0.0104
0.991 ± 0.254
p = 0.0013
0.966 ± 0.108
p < 0.0001
p = 0.4381
S. midas [691-7771]
3304 ± 532
ρ=0.2750
p = 0.3212
ρ=0.5821
p = 0.0228
0.705 ± 0.230
p = 0.0089
0.491 ± 0.268
p = 0.0898
−0.295 ± 0.230
p = 0.2206
0.404 ± 0.194
p = 0.0582
[1987-8209]
4437 ± 550
ρ=0.2912
p = 0.3344
ρ=0.6648
p = 0.0132
1.194 ± 0.389
p = 0.0107
0.715 ± 0.305
p = 0.0390
p = 0.1173
M. nemestrina [1413-8300]
3732 ± 223
ρ=0.0196
p = 0.8839
ρ=0.4969
p = 0.0001
0.469 ± 0.110
p < 0.0001
1.045 ± 0.062
p < 0.0001
−0.531 ± 0.110
p < 0.0001
0.467 ± 0.126
p = 0.0005
[1510-12 511]
4782 ± 295
ρ=0.2114
p = 0.1493
ρ=0.6303
p < 0.0001
1.112 ± 0.172
p < 0.0001
0.902 ± 0.080
p < 0.0001
p = 0.0030
C. apella [106-2901]
1626 ± 96
ρ=0.4959
p = 0.0007
ρ=0.2610
p = 0.0950
0.505 ± 0.106
p < 0.0001
0.633 ± 0.081
p < 0.0001
−0.495 ± 0.106
p < 0.0001
0.282 ± 0.150
p = 0.0684
[271-7371]
2985 ± 244
ρ=0.5126
p = 0.0016
ρ=0.3580
p = 0.0376
0.856 ± 0.164
p < 0.0001
0.400 ± 0.146
p = 0.0100
p < 0.0001

aI, Number of microglial cells; I/mg, number of microglial cells per milligram of structure; N/mg, number of neuronal cells per milligram of structure; O/mg, number of non-neuronal (other) cells per milligram of structure; M, structure mass; I/N, microglia/neuron ratio.

*For Afrotheria, analyses were done, including all the species of the clade (D. dorsalis, E. myurus, L. africana, P. tetradactylus, and P. capensis). Analyses within afrotherian species cortices were only possible where at least three data points were available (L. africana).