Grm4

Models

All models for this gene with cis-heritability P-value <0.01 were tested and are shown here. In each tissue, a gene can have zero, one, or multiple RNA phenotypes for each RNA modality. After the heritability statistics, subsequent columns show cross-validation performance (R2 and P-value) of each modeling method. The best-performing method for each models is used for TWAS.

# Tissue Modality RNA phenotype h2 h2 SE h2 P xQTL R2 BLUP R2 ENET R2 LASSO R2 xQTL P BLUP P ENET P LASSO P
1 NAcc gene expression Grm4 0.18 0.08 9.0e-09 0.04 0.08 0.06 0.08 1.9e-05 2.7e-09 7.4e-08 3.8e-09
2 PL gene expression Grm4 0.05 0.03 7.2e-04 0.04 0.02 0.01 0.03 5.0e-05 5.9e-03 8.8e-03 5.5e-04
3 NAcc mRNA stability Grm4 0.04 0.02 1.2e-05 0.03 0.04 0.03 0.03 2.2e-04 5.3e-05 2.0e-04 1.2e-04
4 Brain mRNA stability Grm4 0.28 0.1 0.0e+00 0.17 0.18 0.19 0.2 7.2e-16 8.9e-17 2.0e-17 2.7e-18

Trait associations

The numbered columns correspond to the numbered models in the table above. They contain the TWAS Z-score for each trait-model pair. Colors indicate strength of Z-scores: cells with lower negative Z-scores are darker blue, cells with higher positive Z-scores are darker red. Significant associations have white and bolded text.

Trait Avg chi2 ratio Avg chi2 Max chi2 1 2 3 4
Bone: apparent density0.50.82.00.3-1.4-0.8-0.7
Area under the delay curve0.30.30.80.60.9-0.10.1
BMI with tail1.55.07.9-2.82.12.61.1
BMI without tail1.12.34.4-0.51.71.32.1
Body weight1.66.713.5-0.23.73.60.6
Weight adjusted by age0.91.02.80.31.71.0-0.2
Bone surface0.40.81.20.9-0.9-0.4-1.1
Bone volume0.20.30.60.6-0.7-0.4-0.5
Breakpoint value in progressive ratio session0.40.41.1-0.3-1.10.5-0.4
Number of licking bursts1.01.22.00.9-1.4-0.8-1.2
Context. condit. distance diff. score6.37.612.32.3-3.5-2.0-2.9
Locomotion velocity, session 11.92.03.31.8-1.1-1.8-0.6
Locomotion distance, session 10.91.23.5-0.3-1.1-1.9-0.2
Locomotion velocity, session 20.91.11.61.3-1.2-1.0-0.7
Locomotion distance, session 20.71.01.50.0-1.1-1.2-1.2
Locomotion velocity, session 30.30.40.70.20.2-0.80.9
Locomotion distance, session 31.31.43.20.4-1.4-1.8-0.3
Stereotopy head waving bouts, day 30.50.61.11.10.50.5-0.9
Stereotopy head waving duration, day 30.30.40.90.90.90.0-0.0
Locomotion velocity, session 72.12.33.20.7-1.8-1.5-1.8
Locomotion distance, session 72.73.45.71.3-2.4-2.3-0.9
Stereotopy head waving bouts, day 70.80.92.70.70.5-0.51.7
Stereotopy head waving duration, day 70.60.71.70.31.00.31.3
Locomotion velocity, session 84.04.78.42.9-2.0-1.3-2.2
Locomotion distance, session 84.87.011.21.4-3.4-2.2-3.1
Degree of sensitization distance0.40.40.60.7-0.50.4-0.8
Degree of sensitization stereotypy1.01.13.10.5-0.1-0.91.8
Conditioned locomotion0.70.82.1-0.2-0.21.5-1.0
Cocaine response after cond. corrected1.01.12.0-1.4-0.40.8-1.3
Cocaine response after cond. not corrected0.30.40.8-0.50.10.7-0.9
Cocaine response before conditioning1.01.22.61.61.2-0.10.7
Saline control response4.76.18.3-2.02.72.32.9
Bone: connectivity density0.91.43.00.3-1.5-0.5-1.7
Bone: cortical apparent density1.72.23.2-1.71.81.21.1
Bone: cortical area0.00.00.0-0.20.20.00.2
Bone: cortical porosity1.71.93.91.20.2-1.52.0
Bone: cortical porosity1.51.63.31.8-0.1-1.31.1
Bone: cortical thickness1.62.65.4-2.31.30.11.8
Bone: cortical thickness3.13.67.2-2.71.61.31.8
Bone: cortical tissue density1.52.03.2-1.51.81.21.1
Condit. Reinf. active minus inactive responses0.40.50.80.9-0.3-0.60.7
Condit. Reinf. active-inactive response ratio0.91.01.9-0.41.10.81.4
Condit. Reinf. active responses0.10.10.20.4-0.2-0.20.3
Condit. Reinf. inactive responses0.10.20.4-0.1-0.6-0.3-0.4
Incentive salience index mean1.51.83.01.5-1.7-1.1-0.9
Condit. Reinf. lever presses3.24.28.01.5-2.8-1.3-2.2
Condit. Reinf. active minus inactive responses0.30.40.50.70.6-0.60.6
Condit. Reinf. active-inactive response ratio0.20.20.40.60.5-0.20.2
Condit. Reinf. active responses0.20.30.50.70.3-0.70.6
Condit. Reinf. inactive responses0.10.10.4-0.10.40.00.6
Condit. Reinf. lever presses0.81.12.00.9-0.5-1.41.1
Condit. Reinf. lever reinforcers received0.40.61.00.80.1-1.00.7
Conditioned reinforcement - actives0.30.30.80.50.4-0.2-0.9
Conditioned reinforcement - inactives0.60.61.3-0.21.0-0.21.2
Conditioned locomotion0.60.81.51.20.0-0.4-1.2
Patch foraging indifference point 0 sec3.15.08.4-1.92.91.72.3
Patch foraging indifference point AUC0.70.82.61.60.4-0.40.4
Delay disc. indifference point, 0s delay0.30.41.20.5-0.2-0.2-1.1
Delay disc. indifference point, 16s delay0.40.40.7-0.40.8-0.40.7
Delay disc. indifference point, 2s delay0.50.61.51.20.7-0.1-0.7
Delay disc. indifference point, 24s delay1.21.22.9-0.21.70.21.3
Delay disc. indifference point, 4s delay0.70.81.71.01.30.7-0.1
Delay disc. indifference point, 8s delay0.20.20.40.60.60.00.3
Indifference point function ln k0.40.61.10.2-1.0-0.9-0.5
Indifference point function log k0.40.61.10.2-1.0-0.9-0.5
Patch foraging total patch changes 0 sec3.55.68.9-2.13.01.72.5
Patch foraging total patch changes 12 sec1.82.44.2-1.42.10.81.6
Patch foraging total patch changes 18 sec3.34.35.0-2.22.21.72.1
Patch foraging total patch changes 24 sec1.72.13.9-1.81.10.42.0
Patch foraging total patch changes 6 sec2.94.26.7-2.02.61.12.3
Patch foraging time to switch 0 sec2.33.59.7-3.10.62.00.2
Patch foraging water rate 0 sec0.20.30.80.30.9-0.2-0.3
Patch foraging water rate 12 sec0.20.40.7-0.4-0.0-0.90.7
Patch foraging water rate 18 sec0.20.20.60.10.80.40.3
Patch foraging water rate 24 sec0.10.10.3-0.10.6-0.30.1
Patch foraging water rate 6 sec0.10.10.4-0.30.3-0.60.2
Seeking ratio, delayed vs. immediate footshock0.60.71.8-0.00.90.61.3
Distance traveled before self-admin0.60.71.7-0.4-0.50.8-1.3
Distance traveled after self-admin0.50.61.7-0.30.71.30.0
Extensor digitorum longus weight0.52.13.71.9-1.0-0.7-1.8
Bone: elastic displacement0.70.81.70.10.7-1.01.3
Bone: elastic work0.30.30.6-0.40.8-0.10.8
Bone: endosteal estimation1.22.54.32.10.42.0-1.4
Bone: endosteal perimeter1.02.13.41.7-1.10.9-1.8
Epididymis fat weight6.816.032.9-5.73.52.33.7
Delta time in closed arm before/after self-admin0.60.61.7-1.30.50.60.1
Delta time in open arm before/after self-admin0.30.31.01.00.3-0.0-0.0
Diff in mean of infusions in LGA sessions1.11.23.1-0.5-1.8-0.5-1.2
Delay discount exponential model param0.30.40.8-0.7-0.90.2-0.2
Extinction: sum of active levers before priming0.30.30.6-0.7-0.8-0.2-0.4
Active lever presses in extinction session 60.40.40.5-0.70.60.60.4
Bone: final force3.25.17.9-2.32.52.81.1
Bone: final moment2.23.55.6-1.62.22.40.9
Food consumed during 24 hour testing period0.81.02.9-0.80.21.7-0.7
Glucose5.07.512.32.9-2.4-1.9-3.5
Heart weight0.50.82.20.70.6-1.50.2
Delay discount hyperbolic model param0.40.40.9-0.9-0.90.2-0.2
Intermittent access intake day 1-15 change0.50.51.30.40.30.61.2
Intermittent access intake escalation0.30.30.50.7-0.20.70.2
Intermittent access intake escalation 23.02.96.92.6-1.6-0.6-1.4
Intermitt. access day 1 inactive lever presses0.60.72.2-1.5-0.5-0.0-0.4
Intermitt. access day 15 inactive lever presses1.11.21.7-0.61.31.11.3
Intermitt. access escalation Index1.31.22.8-0.7-1.7-0.4-1.1
Intermittent access day 1 total infusions0.50.61.30.9-0.3-1.1-0.4
Intermittent access day 15 total infusions0.00.00.10.4-0.0-0.1-0.1
Intermittent access terminal intake (last 3 days)0.10.10.30.60.3-0.30.1
Intermittent access total infusions0.20.20.50.70.4-0.4-0.0
Intermittent access day 1 locomotion0.10.10.20.40.3-0.2-0.4
Intermittent access day 15 total locomotion0.80.81.21.01.01.1-0.3
Intermittent access total locomotion0.80.92.01.40.80.8-0.6
Intermittent access standard deviation2.93.03.8-1.32.01.81.8
Left kidney weight0.51.42.70.41.61.50.8
Right kidney weight0.51.52.81.01.41.70.2
Cocaine induced anxiety1.01.22.30.5-1.50.0-1.5
Post-drug Anxiety0.60.61.7-0.0-0.80.4-1.3
Baseline Anxiety1.11.12.1-1.51.00.41.1
Tail length1.54.718.24.30.7-0.1-0.2
Length with tail1.13.911.93.41.60.8-0.7
Length without tail0.71.52.50.71.61.5-1.0
Times rat made contact with spout0.80.93.0-0.30.4-0.61.7
Lifetime Intake0.10.10.20.50.4-0.3-0.1
Liver weight, left1.11.73.61.90.81.5-0.7
Liver weight, right2.44.013.93.7-0.5-0.3-1.4
Locomotor activity0.30.40.90.30.70.50.9
Locomotor testing distance0.40.71.5-1.20.60.60.7
Locomotor testing rearing0.60.92.1-0.80.71.5-0.4
Locomotion in novel chamber0.10.20.30.60.1-0.5-0.3
Locomotion in novel chamber post-restriction0.91.11.6-1.21.11.30.3
Fecal boli incidents, locomotor time 10.10.10.20.4-0.20.2-0.5
Change in fecal boli incidents, locomotor task0.20.20.7-0.90.10.2-0.3
Time >=10cm from walls, locomotor time 12.22.36.1-1.10.52.5-1.3
Bouts of movement, locomotor time 10.70.81.41.2-1.2-0.4-0.4
Total resting periods, locomotor time 10.80.91.81.3-1.1-0.3-0.7
Rest time, locomotor task time 11.92.15.32.3-0.9-1.0-1.1
Distance moved, locomotor task time 12.12.33.2-1.81.61.51.0
Vertical activity count, locomotor time 14.44.67.4-2.12.32.71.3
Time >=10cm from walls, locomotor time 22.32.45.8-1.31.32.4-0.6
Bouts of movement, locomotor time 20.91.04.02.00.10.20.2
Total resting periods, locomotor time 21.51.65.92.4-0.5-0.3-0.1
Rest time, locomotor task time 22.52.94.81.7-1.9-0.5-2.2
Distance moved, locomotor task time 22.63.04.1-2.02.01.71.2
Vertical activity count, locomotor time 22.22.43.8-1.41.91.80.7
Light reinforcement 11.93.57.0-2.71.12.3-0.1
Light reinforcement 21.01.33.5-0.70.81.9-0.9
Bone: marrow area1.22.64.32.00.52.1-1.3
Bone: maximum diameter1.62.94.82.2-1.6-0.6-2.0
Bone: maximum force1.01.63.0-0.91.71.50.6
Bone: maximum moment0.50.92.1-0.31.51.10.4
Mean time between licks in bursts4.05.78.32.9-2.4-2.0-2.2
Mean num. licks in bursts1.61.93.9-1.01.60.42.0
Cd content in liver1.01.12.0-1.40.81.3-0.5
Co content in liver3.64.27.2-2.71.92.21.1
Cu content in liver0.70.81.5-1.2-0.4-1.10.7
Fe content in liver0.30.41.30.31.10.50.2
K content in liver2.62.63.51.5-1.9-1.8-1.2
Mg content in liver7.37.510.02.8-3.2-2.8-2.1
Mn content in liver4.14.67.02.7-2.0-1.2-2.4
Na content in liver0.10.20.30.2-0.5-0.6-0.0
Rb content in liver0.91.13.20.5-0.90.1-1.8
Se content in liver6.37.712.73.6-2.4-3.2-1.6
Sr content in liver0.30.20.7-0.40.00.4-0.8
Zn content in liver0.70.82.60.4-0.4-0.5-1.6
Bone: minimum diameter0.51.01.71.1-0.50.8-1.3
Time in familiar zone, hab. session 10.40.40.70.3-0.4-0.80.8
Time in novel zone, hab. session 10.40.40.6-0.40.70.6-0.7
Total zone transitions, hab. session 10.81.12.6-0.8-1.0-1.6-0.0
Total locomotion distance, hab. session 11.11.53.6-0.4-1.5-1.9-0.5
Locomotion velocity, hab. session 10.91.23.4-0.3-1.2-1.8-0.2
Time in familiar zone, hab. session 20.10.10.2-0.10.20.10.4
Time in novel zone, hab. session 20.20.20.50.30.70.5-0.3
Total zone transitions, hab. session 20.40.51.1-1.0-0.7-0.6-0.5
Total locomotion distance, hab. session 21.21.62.9-0.9-1.7-1.4-1.0
Locomotion velocity, hab. session 21.72.24.2-0.6-2.1-1.8-1.1
Novel to familiar place preference ratio0.20.20.4-0.5-0.1-0.20.6
Novelty place preference0.10.10.3-0.20.0-0.30.6
Time in novel zone, NPP test0.10.10.20.00.2-0.20.5
Total zone transitions, NPP test1.31.93.5-0.0-1.7-1.9-1.0
Total locomotion distance, NPP test1.11.52.4-0.7-1.5-1.2-1.3
Locomotion velocity, NPP test1.42.03.8-1.1-2.0-1.4-1.0
Intraocular pressure0.20.20.50.5-0.20.3-0.7
Delta distance traveled before/after self-admin1.11.22.40.61.30.51.5
Delta ambulatory time before/after self-admin0.50.60.90.70.80.51.0
Parametrial fat weight0.81.73.5-1.11.21.90.7
Pavlov. Cond. lever latency5.87.310.1-2.83.23.01.5
Pavlov. Cond. magazine entry latency3.84.910.33.2-1.9-2.3-0.6
Pavlov. Cond. change in total contacts1.21.33.11.8-0.9-1.10.0
Pavlov. Cond. index score5.46.810.23.2-2.8-2.8-1.1
Pavlov. Cond. latency score5.36.810.03.2-2.9-2.8-1.1
Pavlov. Cond. lever contacts4.55.27.12.2-2.7-2.6-1.5
Pavlov. Cond. magazine entry number2.73.45.8-2.41.72.10.4
Pavlov. Cond. intertrial magazine entries1.41.73.9-2.01.20.90.7
Pavlov. Cond. lever-magazine prob. diff.5.36.69.73.1-2.7-2.8-1.1
Pavlov. Cond. response bias4.85.99.33.0-2.3-2.7-1.3
Pavlov. Cond. lever latency0.81.02.1-0.90.21.5-1.0
Pavlov. Cond. magazine entry latency0.70.82.61.6-0.7-0.0-0.5
Pavlov. Cond. change in total contacts0.60.92.20.8-0.0-0.91.5
Pavlov. Cond. index score0.60.82.01.4-0.4-0.90.6
Pavlov. Cond. latency score0.50.61.71.3-0.3-0.70.5
Pavlov. Cond. lever contacts0.70.92.70.20.3-0.91.6
Pavlov. Cond. magazine entry number0.10.20.5-0.70.3-0.2-0.0
Pavlov. Cond. intertrial magazine entries0.81.01.71.3-1.2-0.7-0.7
Pavlov. Cond. lever-magazine prob. diff.0.50.71.71.3-0.3-0.70.7
Pavlov. Cond. response bias0.71.01.41.2-0.3-1.21.1
Time in closed arm before self-admin0.10.10.20.5-0.1-0.1-0.5
Time in closed arm after self-admin0.10.10.2-0.50.3-0.20.3
Time in open arm before self-admin0.91.01.91.4-0.7-1.10.7
Time in open arm after self-admin1.21.34.12.0-0.4-0.9-0.0
Bone: periosteal estimation0.40.91.91.00.51.4-0.7
Bone: periosteal perimeter0.20.30.60.7-0.10.7-0.4
Bone: post-yield work0.70.92.01.2-0.40.4-1.4
Progressive ratio test 1 active lever presses0.10.10.30.30.1-0.6-0.1
Progressive ratio test 1 breakpoint0.10.10.3-0.00.2-0.5-0.0
Progressive ratio test 1 inactive lever presses0.91.02.4-1.60.80.80.7
Progressive ratio test 2 active lever presses0.00.00.00.2-0.2-0.2-0.1
Progressive ratio test 2 breakpoint0.10.10.20.2-0.3-0.40.0
Progressive ratio test 2 inactive lever presses0.40.51.51.2-0.1-0.6-0.3
Incentive Sensitization - Responses0.20.20.60.2-0.8-0.3-0.0
Incentive Sensitization - Breakpoint0.20.20.70.2-0.9-0.2-0.1
Sum of active levers in priming session hrs 5-60.20.20.50.2-0.4-0.7-0.1
Progressive ratio4.14.87.0-1.62.62.61.7
Food seeking constrained by brief footshock0.50.62.0-0.10.50.31.4
Seeking ratio, punishment vs. effort1.82.24.30.8-1.9-2.1-0.5
Retroperitoneal fat weight0.20.71.1-0.81.00.90.5
Reaction time number correct1.72.43.6-1.91.41.31.5
Reaction time mean minus median4.05.313.53.7-1.8-0.7-1.9
Reaction time mean minus median AUC4.05.212.73.6-1.7-0.8-2.1
Reaction time num false alarms0.20.30.50.7-0.6-0.4-0.0
Reaction time num false alarms AUC0.20.30.40.7-0.5-0.60.2
Reaction time trials correct on left1.72.43.6-1.91.41.31.5
Reaction time trials on left1.62.33.3-1.81.41.31.5
Reaction time mean2.63.65.92.4-1.7-0.0-2.4
Reaction time mean AUC2.53.46.42.5-1.60.1-2.2
Median of all reaction times1.41.94.11.3-1.30.3-2.0
Reaction time omissions1.62.23.71.9-1.6-0.1-1.6
Reaction time false alarm rate1.31.82.91.7-1.2-1.5-0.6
Reaction time premature initiation rate0.10.20.4-0.70.30.40.0
Reaction time premature initiations1.21.83.0-1.71.31.30.9
Std. dev. reaction times3.65.113.63.7-1.7-0.6-1.9
Reaction time trials completed1.62.33.3-1.81.41.31.5
Reaction time trials AUC1.62.43.7-1.91.41.41.4
Run reversals in cocaine runway, females0.60.61.0-0.70.90.11.0
Run reversals in cocaine runway, males2.72.66.12.5-1.4-0.9-1.2
Latency to leave start box in cocaine runway0.50.61.2-0.8-1.1-0.5-0.4
Latency to leave start box in cocaine runway, F0.40.41.0-0.5-1.0-0.4-0.6
Latency to leave start box in cocaine runway, M0.40.31.1-1.0-0.3-0.30.4
Std. dev. time between licks in bursts4.04.85.32.2-2.3-1.9-2.2
Total sessions with >9 infusions0.20.20.30.6-0.2-0.5-0.6
Short access day 1 total inactive lever presses1.41.51.81.3-1.3-1.0-1.3
Short access day 10 total inactive lever presses0.50.51.0-1.0-0.90.40.3
Short access day 1 total infusions0.70.81.90.9-0.5-1.4-0.3
Short access day 10 total infusions0.10.10.2-0.20.50.20.1
Short access total infusions0.10.10.30.30.3-0.50.1
Short access day 1 locomotion0.70.71.7-0.5-1.3-0.8-0.4
Short access day 10 total locomotion0.20.20.90.3-0.1-0.1-0.9
Short access total locomotion0.10.10.2-0.40.10.2-0.5
One hour access (0.3 mA shock)1.21.42.20.8-1.5-1.1-1.3
Compulsive drug intake1.71.74.2-0.3-2.1-0.8-1.3
One hour access (shock baseline)0.81.03.41.80.5-0.2-0.4
Social response proportion0.10.10.3-0.10.60.3-0.4
Social responses0.10.10.2-0.30.40.30.2
Social time2.12.74.51.6-2.1-0.5-1.9
Soleus weight2.05.910.93.1-1.5-0.8-3.3
Bone: stiffness0.91.42.3-1.21.31.50.5
Tibialis anterior weight0.00.20.3-0.5-0.0-0.40.4
Time to tail flick, vehicle, before self-admin0.50.50.8-0.3-0.80.6-0.9
Time to tail flick, vehicle, after self-admin0.40.40.70.4-0.70.4-0.8
Time to tail flick, test, before self-admin0.30.30.7-0.50.2-0.20.8
Time to tail flick, test, after self-admin2.12.24.22.0-1.6-1.3-0.1
Delta time to tail flick, vehicle, before/after SA0.20.20.40.5-0.2-0.60.3
Delta time to tail flick, test, before/after SA0.30.30.90.2-1.0-0.1-0.3
Tibia length0.61.73.41.30.41.9-1.2
Ambulatory time before self-admin0.30.30.5-0.4-0.70.3-0.7
Ambulatory time after self-admin0.20.20.60.10.30.80.1
Bone: tissue strength0.30.30.6-0.30.80.7-0.3
Total heroin consumption0.00.00.10.1-0.30.10.1
Bone: trabecular number0.60.92.10.1-1.5-1.0-0.6
Bone: trabecular spacing1.31.83.3-0.41.81.61.0
Bone: trabecular thickness0.70.81.80.9-1.3-0.3-0.8
Bone: trabecular tissue density0.71.02.61.60.2-0.71.0
Water consumed over 24 hours0.20.30.70.1-0.5-0.1-0.8
Number of responses in last shaping day1.41.54.41.0-0.5-2.1-0.3