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Edited by Huda Akil, University of Michigan, Ann Arbor, MI, and approved December 16, 2019 (received for review June 10, 2019)
Article Figures & SI
Figures
- Fig. 1.
Experimental design. The mice underwent the 2BC/CIE paradigm. The 2BC/CIE paradigm involves alternating weeks of 2BC (highlighted in blue) and CIE/Air (highlighted in red and gray, respectively). The mice underwent 2 wk of baseline 2BC testing followed by five rounds of alternating CIE or Air weeks and 2BC weeks. The mice then underwent a sixth week of CIE/Air and were then tested for irritability-like and digging behaviors 7 and 10 d, respectively, into abstinence (highlighted in green). The mice then underwent one additional week of CIE/Air, and brains were collected 7 d after the last vapor exposure with no intervening behavioral testing. Brains were collected during the time of day that the mice would normally undergo a 2BC session. Brains were then immunostained for Fos and cleared using the iDISCO+ procedure. Brains were then imaged and analyzed to identify brain regions that potentially contribute to behaviors and the functional networks for each treatment. Key brain regions in the alcohol abstinence network were then identified using graph theory. BBT, bottle-brush test of irritability-like behavior; DM, digging and marble burying.
- Fig. 2.
Alcohol drinking and abstinence behavior in alcohol-dependent vs. control mice. (A) Alcohol-dependent mice (black bars) exhibited a significant increase in alcohol drinking during postvapor week-5 2BC testing compared with alcohol drinking in nondependent mice (white bars) and their own baseline intake. (B) Irritability-like behavior. Alcohol-dependent mice (black bar) exhibited a significant increase in total irritable-like responses compared with control mice (nondependent and naive; white bar). (C) Digging behavior. Alcohol-dependent mice (black bar) exhibited a significant increase in composite digging behavior (Z-score of digging behaviors) compared with control mice (nondependent and naive; white bar). *P < 0.05 (two-tailed), alcohol-dependent vs. nondependent for alcohol drinking or control for irritability-like and digging behavior; #P < 0.05 (two-tailed), alcohol-dependent postvapor week 5 vs. alcohol-dependent baseline.
- Fig. 3.
Interbrain regional Pearson correlation heat maps for each treatment organized anatomically based on the Allen Mouse Brain Atlas. (A) Correlation heat map for naive mice. (B) Correlation heat map for alcohol-nondependent mice. (C) Correlation heat map for alcohol-dependent mice. Each heat map is organized into color-coded anatomical groups: dark green (cortical plate), light green (cortical subplate), dark blue (striatum), light blue (pallidum), dark red (thalamus), light red (hypothalamus), and purple (midbrain, hindbrain, and cerebellum). The region that is highlighted in purple on each heat map represents an amygdala cluster of the heat map that is shown in greater detail in Fig. 4.
- Fig. 4.
Comparison of Pearson correlations of an amygdala cluster vs. other regions. (A) Cutout of correlations from Fig. 3 of brain regions compared with the amygdala cluster for each treatment. Individual region names are displayed at the bottom, and group names are displayed above each cluster. (B) Average R values for alcohol-dependent (black bars), alcohol-nondependent (white bars), and naive (gray bars) mice for each cluster vs. the amygdala cluster. *P < 0.05, vs. naive; #P < 0.05, vs. alcohol-nondependent.
- Fig. 5.
Hierarchical clustering of complete Euclidean distance matrices for each treatment. Modules were determined by cutting each dendrogram at half of the maximal tree height. (A) Relative distance of each brain region relative to the others that were examined in naive mice. In naive mice, nine distinct modules of coactivation were identified. (B) Relative distance of each brain region relative to the others that were examined in alcohol-nondependent mice. In alcohol-nondependent mice, eight distinct modules of coactivation were identified. (C) Relative distance of each brain region relative to the others that were examined in alcohol-dependent mice. In alcohol-dependent mice, three distinct modules of coactivation were identified. For all distance matrices, each module is boxed in purple. (D) Number of modules in each treatment condition after cutting the hierarchical clustered dendrogram at different percentages of tree height. In all cases (except at extreme cutoff values; e.g., 70–100%), the alcohol-dependent network showed a lower number of modules compared with the alcohol-nondependent and naive networks.
- Fig. 6.
Functional connectivity of alcohol-dependent mice during abstinence thresholded to 0.75R. Nodes/brain regions of the network are represented by circles. The size of the node represents the PC (smaller = lower PC; larger = higher PC). The internal color of each circle represents the within-module degree Z-score (dark blue = lowest; dark red = highest). The color of the modules that are identified in Fig. 5C are represented by different colored edges of each node circle (red = module A/extended amygdala; blue = module B/midbrain striatal; green = module C/cortico-hippocampo-thalamic). The thickness of the lines represents the strength of the correlation between regions (thin = lower correlation; thick = higher correlation). See figure key for examples of each representative component of the figure.
- Fig. 7.
Theories of brain regions that are involved in the neurobiology of alcohol use disorder. (A) Brain regions of the mesolimbic dopamine system. (B) Brain regions consisting of cortico-striatal loops. (C) Extended amygdalar brain regions. (D) Three-stage theory. Images are modified from ref. 4.
Data supplements
Supporting Information
Brain-wide functional architecture remodeling by alcohol dependence and abstinence
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