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. 2018 Feb 21;24(9):1329–1350. doi: 10.1038/s41380-018-0025-5

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© The Author(s) 2018

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 3 — Taok2 KO mice have reduced dendrite growth and synaptic connectivity in the prefrontal cortex. a Top: Golgi-stained PFC neurons from P21 WT, Taok2 Het and KO mice. Scale bars represent 20 μm. Bottom: dendritic heat maps of superimposed neuron tracings for each condition. Blue to red (apical) and yellow to blue (basal) indicates increased probability of dendrite presence. Scale bars represent 30 μm. b Top: no major difference in apical dendritic complexity in layer 2 PFC neurons in Taok2 Het and KO mice (WT = 26, Het = 32, KO = 21 neurons from three different brains; two-way ANOVA, post hoc Dunnett’s test; F_2, 3496 = 3.055, p = 0.0472 between genotypes; *represents ranges of significance; WT vs. Het (blue), WT vs. KO (red); see supplemental statistics). Bottom: significantly reduced basal dendritic complexity in layer 2 PFC neurons in Taok2 Het and KO mice (WT = 19, Het = 18 and KO = 18 neurons from three different brains; two-way ANOVA, post hoc Dunnett’s test; F_2, 1716 = 128.7, p < 0.0001 between genotypes; *represents ranges of significance; WT vs. Het (blue), WT vs. KO (red); see supplemental statistics). c Top: no change in apical dendrite length (µm) in Taok2 KO PFC neurons (WT = 27, Het = 34, and KO = 22 neurons from three different mice brains; one-way ANOVA, post hoc Dunnet’s test; F_{2, 80} = 0.3346, p = 0.7167). Bottom: reduced basal dendrite length (µm) in Taok2 KO PFC neurons (WT = 22, Het = 17, and KO = 18 neurons from three different mice brains; one-way ANOVA, post hoc Dunnet’s test; F_{2, 54} = 12.47, p < 0.0001; WT vs. Het p = 0.0007, WT vs. KO p = 0.0001). d Dendritic spine distribution maps of Golgi-stained WT and Taok2 KO PFC neurons. e Taok2 KO PFC neurons show reduced number of distal dendritic spines (WT = 6, Het = 6, and KO = 6 neurons from three different brains; two-way ANOVA, post hoc Dunnett’s test; F_{2, 465} = 89.35, p < 0.0001 between genotypes; *represents ranges of significance; WT vs. Het (blue), WT vs. KO (red); see supplemental statistics). f Left: Taok2 KO PFC neurons show no difference in total apical dendritic spines per cell (WT = 6, Het = 6, KO = 6 neurons from three different mice brains; One-way ANOVA, post hoc Dunnett’s test; F_2, 15 = 1.766, p = 0.2048). Right: Taok2 KO PFC neurons show decreased number of total basal dendritic spines per cell (WT = 6, Het = 6, KO = 6 neurons from three different mice brains; One-way ANOVA, post hoc Dunnett’s test; F_{2, 15} = 13.76, p = 0.0004; WT vs. Het p = 0.0265, WT vs. KO p = 0.0002. g Images of dendritic spines on P21 WT and Taok2 KO PFC neuron dendrites. h Cumulative probability histograms show shift toward reduced dendritic spine lengths (left) Taok2 KO PFC neurons and reduced head widths (right) in Taok2 Het and KO PFC neurons. i Taok2 KO PFC neurons have a significant increase in thin and stubby shaped spines and reduction in mushroom-like spines compared with WT PFC neurons, whereas Taok2 Het PFC neurons have an increase in long thin shaped spines and reduction in mushroom-like spines (WT = 3504, Het = 2317, KO = 1262 spines from six cells per condition from three different brains; two-way ANOVA, post hoc Dunnett’s test; F_2, 90 = 0.0008294, p = 0.9992 between genotypes; WT vs. Het filopodia: p = 0.9997, long thin: p = 0.0088, thin: p = 0.5599, stubby: p = 0.8934, mushroom: p < 0.0001, and branched: p = 0.9765; WT vs. KO filopodia: p = 0.5111, long thin: p = 0.7565, thin: p = 0.0141, stubby: p < 0.0001, mushroom: p < 0.0001, and branched: p = 0.9943). j Representative images of synapses innervating spines (top) and synapses innervating dendrites (bottom) and illustrations of each (right). k Taok2 KO neurons imaged by electron microscopy show decreased percentage of synapses on postsynaptic spines (left) and increased synapses on dendrites (right) (WT = 159 and KO = 120 neurons; innervating spine: unpaired t-test; t(277) = 8.814, p < 0.0001; innervating dendrite: t(277) = 8.262, p < 0.0001). l Representative traces of mEPSC spikes from WT and Taok2 KO PFC neurons. Scale: 5pA vs. 1 s. m Longer inter-event intervals in Taok2 Het and KO PFC neurons shown on a cumulative probability histogram. Inside: reduced mean mEPSC event frequency in Taok2 Het and KO PFC neurons (WT = 14, Het = 13 and KO = 13 neurons from three different mice brains; Kruskal-Walis ANOVA, post hoc Dunn’s test; H-value = 14.47, p = 0.0007; WT vs. Het p = 0.0212, WT vs. KO p = 0.0008). n No change in mEPSC amplitude in Taok2 WT, Het and KO PFC neurons shown on a cumulative probability histogram. Inside: mean mEPSC amplitude of Taok2 WT, Het and KO neurons (WT = 14, Het = 13 and KO = 13 neurons from three different mice brains; kruskal-walis ANOVA, post hoc Dunn’s test; H-value = 3.871, p = 0.1444; WT vs. Het p = 0.1857, WT vs. KO p > 0.9999). *p < 0.05, **p < 0.01, ***p < 0.001 and ****p < 0.0001. Values are mean ± s.e.m.

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