Ell

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 Ell 0.04 0.03 2.1e-06 0.05 0.04 0.05 0.05 1.8e-08 3.6e-07 1.2e-07 3.3e-08
2 BLA gene expression Ell 0.09 0.05 4.7e-04 0.05 0.05 0.02 0.03 1.3e-03 1.5e-03 3.3e-02 8.8e-03
3 Adipose mRNA stability Ell 0.11 0.05 1.4e-14 0.08 0.12 0.13 0.12 2.7e-09 1.1e-13 3.1e-14 9.0e-14
4 Brain mRNA stability Ell 0.05 0.03 6.1e-04 0.05 0.03 0.04 0.04 1.4e-05 1.6e-03 1.7e-04 1.2e-04

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 density0.40.61.5-0.7-0.4-0.61.2
Area under the delay curve0.00.10.10.30.3-0.00.1
BMI with tail1.96.59.4-2.4-2.4-3.1-2.3
BMI without tail3.410.414.3-3.1-3.1-3.8-2.8
Body weight4.921.329.2-5.0-4.7-5.4-3.0
Weight adjusted by age1.52.33.1-1.7-1.7-1.7-0.6
Bone surface0.10.20.6-0.5-0.2-0.8-0.1
Bone volume0.50.81.6-1.1-0.8-1.30.0
Breakpoint value in progressive ratio session0.10.10.3-0.1-0.20.1-0.6
Number of licking bursts0.10.20.50.40.40.10.7
Context. condit. distance diff. score0.20.20.60.20.10.50.8
Locomotion velocity, session 10.60.61.60.70.7-0.3-1.3
Locomotion distance, session 11.41.83.21.71.81.10.3
Locomotion velocity, session 21.41.62.1-1.1-1.1-1.5-1.3
Locomotion distance, session 21.92.74.41.92.11.60.5
Locomotion velocity, session 30.60.61.20.80.61.10.6
Locomotion distance, session 30.80.90.90.91.00.90.9
Stereotopy head waving bouts, day 31.82.14.10.80.61.82.0
Stereotopy head waving duration, day 30.50.61.30.30.11.11.1
Locomotion velocity, session 71.01.11.61.21.31.1-0.1
Locomotion distance, session 70.91.21.91.31.41.00.3
Stereotopy head waving bouts, day 70.30.30.9-0.2-0.20.60.9
Stereotopy head waving duration, day 70.30.30.7-0.7-0.90.10.3
Locomotion velocity, session 81.31.62.51.61.31.4-0.1
Locomotion distance, session 80.71.11.81.31.30.90.1
Degree of sensitization distance0.10.10.10.40.30.2-0.2
Degree of sensitization stereotypy0.20.20.3-0.6-0.6-0.10.4
Conditioned locomotion0.00.00.2-0.00.00.10.4
Cocaine response after cond. corrected0.40.51.2-0.3-0.40.71.1
Cocaine response after cond. not corrected0.40.51.10.20.11.00.9
Cocaine response before conditioning0.60.71.21.11.10.4-0.6
Saline control response0.30.30.60.60.60.4-0.8
Bone: connectivity density0.20.30.9-0.5-0.3-0.90.2
Bone: cortical apparent density0.20.30.50.60.70.10.5
Bone: cortical area4.67.110.4-3.2-3.0-2.6-1.3
Bone: cortical porosity0.20.20.70.30.4-0.2-0.9
Bone: cortical porosity0.20.20.60.30.4-0.3-0.7
Bone: cortical thickness2.33.96.6-2.6-2.4-1.70.3
Bone: cortical thickness1.41.63.9-1.2-1.1-0.12.0
Bone: cortical tissue density0.30.40.70.70.80.20.5
Condit. Reinf. active minus inactive responses0.30.40.9-0.6-0.4-0.11.0
Condit. Reinf. active-inactive response ratio0.50.51.8-0.5-0.30.01.3
Condit. Reinf. active responses0.30.40.6-0.8-0.7-0.50.2
Condit. Reinf. inactive responses0.91.12.1-0.7-0.8-1.0-1.5
Incentive salience index mean0.30.30.6-0.8-0.6-0.7-0.0
Condit. Reinf. lever presses0.50.71.0-0.8-0.6-1.0-1.0
Condit. Reinf. active minus inactive responses0.20.20.30.60.50.60.3
Condit. Reinf. active-inactive response ratio0.30.40.5-0.6-0.7-0.30.7
Condit. Reinf. active responses0.91.22.01.41.31.0-0.3
Condit. Reinf. inactive responses2.02.63.82.01.91.2-1.3
Condit. Reinf. lever presses0.91.24.5-0.1-0.2-0.5-2.1
Condit. Reinf. lever reinforcers received0.40.50.90.70.60.4-0.9
Conditioned reinforcement - actives1.22.13.6-1.4-1.6-1.9-0.1
Conditioned reinforcement - inactives6.19.014.0-3.7-3.6-3.00.0
Conditioned locomotion1.41.83.4-1.3-1.4-1.80.2
Patch foraging indifference point 0 sec2.34.34.9-2.1-2.1-2.2-1.8
Patch foraging indifference point AUC0.20.31.10.20.2-0.2-1.0
Delay disc. indifference point, 0s delay2.22.44.02.01.81.40.8
Delay disc. indifference point, 16s delay0.40.40.80.90.80.40.1
Delay disc. indifference point, 2s delay0.70.81.31.01.10.80.6
Delay disc. indifference point, 24s delay0.40.40.7-0.3-0.7-0.8-0.7
Delay disc. indifference point, 4s delay0.00.10.10.30.3-0.1-0.0
Delay disc. indifference point, 8s delay0.10.10.2-0.3-0.4-0.4-0.3
Indifference point function ln k0.50.91.4-1.1-1.0-1.20.0
Indifference point function log k0.50.91.4-1.1-1.0-1.20.0
Patch foraging total patch changes 0 sec3.56.77.7-2.7-2.7-2.8-2.1
Patch foraging total patch changes 12 sec0.60.81.1-0.9-0.9-0.9-1.0
Patch foraging total patch changes 18 sec0.10.10.40.0-0.00.2-0.6
Patch foraging total patch changes 24 sec0.20.30.5-0.7-0.7-0.4-0.1
Patch foraging total patch changes 6 sec1.42.13.0-1.2-1.3-1.5-1.7
Patch foraging time to switch 0 sec0.81.72.51.51.61.40.2
Patch foraging water rate 0 sec0.91.92.8-1.4-1.4-1.7-0.9
Patch foraging water rate 12 sec0.40.71.7-0.8-0.8-1.3-0.2
Patch foraging water rate 18 sec0.10.20.40.40.4-0.7-0.1
Patch foraging water rate 24 sec0.30.41.4-0.4-0.3-1.2-0.1
Patch foraging water rate 6 sec1.52.23.5-1.5-1.6-1.9-0.5
Seeking ratio, delayed vs. immediate footshock0.50.61.50.81.20.4-0.2
Distance traveled before self-admin0.70.91.0-0.9-1.0-1.0-0.9
Distance traveled after self-admin1.11.41.6-1.3-1.3-1.0-1.1
Extensor digitorum longus weight0.20.70.9-0.9-0.9-0.9-0.6
Bone: elastic displacement0.40.40.60.70.80.50.6
Bone: elastic work1.11.42.2-1.5-1.4-1.3-0.3
Bone: endosteal estimation0.81.84.9-0.6-0.5-1.2-2.2
Bone: endosteal perimeter1.32.83.2-1.8-1.8-1.6-1.4
Epididymis fat weight1.44.67.7-1.5-1.2-2.6-2.8
Delta time in closed arm before/after self-admin0.30.20.5-0.2-0.2-0.6-0.7
Delta time in open arm before/after self-admin0.70.61.00.90.91.0-0.0
Diff in mean of infusions in LGA sessions1.41.71.8-1.3-1.3-1.3-1.3
Delay discount exponential model param0.00.00.0-0.0-0.00.2-0.0
Extinction: sum of active levers before priming0.50.61.10.91.10.5-0.5
Active lever presses in extinction session 60.30.30.40.70.40.4-0.6
Bone: final force6.710.613.5-3.7-3.4-3.5-2.2
Bone: final moment6.510.413.2-3.6-3.4-3.5-2.2
Food consumed during 24 hour testing period0.40.61.10.91.10.60.0
Glucose1.72.63.1-1.5-1.5-1.7-1.7
Heart weight1.01.62.2-1.3-1.4-0.61.5
Delay discount hyperbolic model param0.00.00.0-0.1-0.10.1-0.1
Intermittent access intake day 1-15 change0.10.20.3-0.6-0.5-0.20.5
Intermittent access intake escalation0.10.20.5-0.3-0.10.10.7
Intermittent access intake escalation 20.40.50.6-0.7-0.7-0.8-0.7
Intermitt. access day 1 inactive lever presses0.10.20.5-0.4-0.7-0.30.4
Intermitt. access day 15 inactive lever presses0.10.20.8-0.3-0.30.30.9
Intermitt. access escalation Index0.30.50.60.50.80.60.8
Intermittent access day 1 total infusions0.91.42.1-1.4-1.4-1.3-0.4
Intermittent access day 15 total infusions0.40.61.1-1.1-0.8-0.50.7
Intermittent access terminal intake (last 3 days)0.40.61.0-1.0-0.9-0.60.2
Intermittent access total infusions1.41.62.8-1.7-1.5-1.2-0.2
Intermittent access day 1 locomotion2.12.74.3-1.6-1.8-2.1-0.7
Intermittent access day 15 total locomotion0.20.30.5-0.3-0.7-0.70.1
Intermittent access total locomotion1.82.23.4-1.5-1.7-1.8-0.4
Intermittent access standard deviation0.20.40.6-0.5-0.7-0.8-0.3
Left kidney weight0.62.95.2-1.4-1.4-1.7-2.3
Right kidney weight1.98.614.1-2.4-2.3-3.1-3.8
Cocaine induced anxiety1.11.62.1-1.4-1.4-1.1-1.1
Post-drug Anxiety0.10.10.20.0-0.2-0.0-0.5
Baseline Anxiety2.63.04.52.11.91.80.8
Tail length0.93.64.8-1.9-1.8-2.2-1.7
Length with tail1.88.010.0-3.2-2.9-3.1-2.0
Length without tail1.13.66.3-2.5-2.3-1.7-0.1
Times rat made contact with spout0.40.51.20.50.60.21.1
Lifetime Intake2.53.04.8-2.2-2.0-1.7-0.1
Liver weight, left0.40.71.30.91.10.8-0.1
Liver weight, right2.13.76.3-1.5-1.3-2.1-2.5
Locomotor activity0.10.10.20.10.5-0.1-0.4
Locomotor testing distance0.30.50.90.70.90.50.5
Locomotor testing rearing0.20.51.00.91.00.2-0.2
Locomotion in novel chamber0.10.10.30.00.1-0.5-0.5
Locomotion in novel chamber post-restriction0.10.10.20.40.3-0.30.2
Fecal boli incidents, locomotor time 10.10.10.30.60.4-0.0-0.0
Change in fecal boli incidents, locomotor task0.20.20.40.50.10.6-0.4
Time >=10cm from walls, locomotor time 10.20.20.6-0.0-0.1-0.3-0.7
Bouts of movement, locomotor time 10.00.00.10.30.2-0.1-0.2
Total resting periods, locomotor time 10.20.20.40.60.60.20.0
Rest time, locomotor task time 10.00.00.1-0.0-0.3-0.0-0.2
Distance moved, locomotor task time 10.00.10.10.20.20.40.1
Vertical activity count, locomotor time 10.00.10.20.10.40.2-0.1
Time >=10cm from walls, locomotor time 21.11.61.91.21.21.41.2
Bouts of movement, locomotor time 20.10.20.40.2-0.10.60.6
Total resting periods, locomotor time 20.10.20.3-0.1-0.30.50.5
Rest time, locomotor task time 20.70.92.2-1.1-1.5-0.5-0.3
Distance moved, locomotor task time 20.40.50.70.70.90.70.5
Vertical activity count, locomotor time 20.50.71.40.40.61.01.2
Light reinforcement 10.10.30.7-0.1-0.1-0.6-0.8
Light reinforcement 20.30.51.4-0.5-0.5-0.4-1.2
Bone: marrow area0.71.54.4-0.5-0.4-1.1-2.1
Bone: maximum diameter2.24.16.3-2.5-2.5-1.8-0.8
Bone: maximum force5.69.912.3-3.5-3.2-3.5-2.2
Bone: maximum moment5.08.811.1-3.3-3.0-3.2-2.1
Mean time between licks in bursts1.53.85.3-2.0-1.9-2.3-1.4
Mean num. licks in bursts0.10.10.3-0.2-0.10.10.5
Cd content in liver1.31.42.4-1.5-1.5-0.80.7
Co content in liver0.10.20.5-0.00.10.30.7
Cu content in liver2.02.33.31.31.41.51.8
Fe content in liver0.70.81.30.90.91.10.6
K content in liver2.01.92.51.51.51.60.8
Mg content in liver0.10.10.30.30.30.6-0.0
Mn content in liver0.20.20.8-0.2-0.1-0.0-0.9
Na content in liver2.72.94.72.22.21.4-0.4
Rb content in liver1.31.52.5-1.5-1.6-0.41.1
Se content in liver0.50.60.90.70.80.90.7
Sr content in liver0.20.20.40.40.6-0.1-0.6
Zn content in liver0.00.00.10.0-0.10.1-0.2
Bone: minimum diameter2.74.96.0-2.1-1.9-2.4-2.4
Time in familiar zone, hab. session 10.30.30.8-0.3-0.4-0.9-0.4
Time in novel zone, hab. session 10.20.20.60.00.10.40.8
Total zone transitions, hab. session 10.50.71.11.01.00.1-1.0
Total locomotion distance, hab. session 11.01.42.71.51.70.6-0.5
Locomotion velocity, hab. session 11.01.42.61.51.60.4-0.8
Time in familiar zone, hab. session 20.30.30.6-0.2-0.3-0.6-0.8
Time in novel zone, hab. session 20.30.30.9-0.10.00.40.9
Total zone transitions, hab. session 20.10.10.20.40.50.40.0
Total locomotion distance, hab. session 20.40.60.70.70.80.60.8
Locomotion velocity, hab. session 20.50.61.00.81.00.60.6
Novel to familiar place preference ratio0.20.30.4-0.5-0.5-0.4-0.6
Novelty place preference0.10.10.2-0.4-0.4-0.2-0.4
Time in novel zone, NPP test0.00.00.1-0.1-0.1-0.0-0.4
Total zone transitions, NPP test0.71.12.11.31.40.70.3
Total locomotion distance, NPP test1.62.23.81.72.01.00.8
Locomotion velocity, NPP test1.42.03.61.71.91.00.6
Intraocular pressure1.11.63.0-1.7-1.6-0.60.6
Delta distance traveled before/after self-admin0.60.71.4-1.2-0.8-0.6-0.5
Delta ambulatory time before/after self-admin0.10.10.3-0.5-0.5-0.00.2
Parametrial fat weight4.310.112.2-3.1-3.0-3.5-3.2
Pavlov. Cond. lever latency0.00.00.1-0.1-0.30.2-0.2
Pavlov. Cond. magazine entry latency0.20.20.6-0.4-0.3-0.8-0.1
Pavlov. Cond. change in total contacts0.70.71.0-1.0-0.8-1.00.3
Pavlov. Cond. index score0.10.10.10.10.3-0.4-0.1
Pavlov. Cond. latency score0.10.10.30.30.5-0.20.1
Pavlov. Cond. lever contacts0.10.10.3-0.20.0-0.5-0.2
Pavlov. Cond. magazine entry number0.30.30.90.40.31.00.3
Pavlov. Cond. intertrial magazine entries1.21.44.40.30.31.02.1
Pavlov. Cond. lever-magazine prob. diff.0.00.10.10.10.2-0.4-0.1
Pavlov. Cond. response bias0.20.20.6-0.3-0.2-0.8-0.0
Pavlov. Cond. lever latency1.41.95.40.91.00.82.3
Pavlov. Cond. magazine entry latency0.50.61.0-0.7-0.8-0.5-1.0
Pavlov. Cond. change in total contacts0.91.43.9-0.8-0.9-0.6-2.0
Pavlov. Cond. index score1.41.83.9-1.1-1.2-0.9-2.0
Pavlov. Cond. latency score0.81.02.7-0.7-0.8-0.5-1.7
Pavlov. Cond. lever contacts1.11.54.4-0.7-0.9-0.6-2.1
Pavlov. Cond. magazine entry number1.92.34.61.31.41.22.1
Pavlov. Cond. intertrial magazine entries0.40.61.21.11.10.30.3
Pavlov. Cond. lever-magazine prob. diff.1.51.94.0-1.1-1.3-1.0-2.0
Pavlov. Cond. response bias1.72.46.0-1.0-1.1-1.0-2.5
Time in closed arm before self-admin4.55.89.3-3.1-2.9-2.1-0.9
Time in closed arm after self-admin4.34.57.1-2.7-2.5-2.1-0.6
Time in open arm before self-admin0.80.91.91.21.40.50.2
Time in open arm after self-admin2.22.54.01.92.01.40.5
Bone: periosteal estimation2.14.55.0-2.2-2.0-2.2-2.0
Bone: periosteal perimeter4.17.010.3-3.2-3.2-2.5-1.2
Bone: post-yield work0.00.00.00.10.20.1-0.2
Progressive ratio test 1 active lever presses2.63.24.6-2.0-2.0-2.2-0.4
Progressive ratio test 1 breakpoint2.42.94.2-2.0-1.9-2.1-0.4
Progressive ratio test 1 inactive lever presses1.21.72.4-1.5-1.6-1.30.6
Progressive ratio test 2 active lever presses0.91.11.7-1.3-1.3-1.0-0.0
Progressive ratio test 2 breakpoint0.70.81.2-1.1-1.1-0.90.1
Progressive ratio test 2 inactive lever presses1.21.93.1-1.8-1.7-1.10.8
Incentive Sensitization - Responses0.10.10.2-0.5-0.4-0.00.3
Incentive Sensitization - Breakpoint0.00.00.1-0.2-0.10.10.3
Sum of active levers in priming session hrs 5-64.23.94.62.02.22.11.6
Progressive ratio0.10.10.30.10.00.4-0.5
Food seeking constrained by brief footshock0.80.92.1-0.10.51.21.5
Seeking ratio, punishment vs. effort0.60.92.70.40.70.51.6
Retroperitoneal fat weight2.912.216.5-3.6-3.5-4.1-2.7
Reaction time number correct0.20.30.40.60.7-0.5-0.5
Reaction time mean minus median0.10.20.70.1-0.10.0-0.8
Reaction time mean minus median AUC0.10.20.5-0.2-0.4-0.1-0.7
Reaction time num false alarms0.60.81.8-1.2-1.3-0.3-0.4
Reaction time num false alarms AUC0.40.61.4-1.0-1.20.1-0.3
Reaction time trials correct on left0.20.30.40.60.7-0.5-0.5
Reaction time trials on left0.20.40.50.50.6-0.6-0.7
Reaction time mean0.91.51.9-1.2-1.4-0.9-1.4
Reaction time mean AUC0.60.92.0-0.7-0.9-0.6-1.4
Median of all reaction times1.01.42.1-1.3-1.5-0.9-1.1
Reaction time omissions2.13.14.9-1.5-1.7-1.5-2.2
Reaction time false alarm rate0.70.92.0-1.3-1.40.00.1
Reaction time premature initiation rate0.30.40.60.70.70.50.8
Reaction time premature initiations0.10.20.50.70.7-0.10.1
Std. dev. reaction times0.61.12.2-0.8-0.9-0.8-1.5
Reaction time trials completed0.20.40.50.50.6-0.6-0.7
Reaction time trials AUC0.20.40.70.30.4-0.8-0.8
Run reversals in cocaine runway, females4.54.57.0-2.7-2.4-2.0-1.1
Run reversals in cocaine runway, males1.11.11.31.11.11.10.8
Latency to leave start box in cocaine runway0.30.30.7-0.1-0.1-0.7-0.8
Latency to leave start box in cocaine runway, F0.50.61.0-0.8-0.6-1.0-0.6
Latency to leave start box in cocaine runway, M0.30.30.50.70.50.3-0.5
Std. dev. time between licks in bursts1.92.53.7-1.7-1.6-1.9-0.9
Total sessions with >9 infusions4.45.47.7-2.6-2.6-2.8-0.3
Short access day 1 total inactive lever presses1.72.33.3-1.6-1.8-1.51.0
Short access day 10 total inactive lever presses0.10.30.60.60.20.80.1
Short access day 1 total infusions3.64.76.8-2.5-2.4-2.60.4
Short access day 10 total infusions1.31.62.7-1.5-1.3-1.70.1
Short access total infusions2.93.65.2-2.2-2.1-2.3-0.1
Short access day 1 locomotion5.56.89.7-3.1-3.0-2.8-0.7
Short access day 10 total locomotion1.22.23.6-1.7-1.5-1.90.1
Short access total locomotion2.93.44.8-2.1-2.0-2.2-0.4
One hour access (0.3 mA shock)0.70.81.4-1.0-0.9-0.31.2
Compulsive drug intake0.91.02.20.80.81.50.8
One hour access (shock baseline)1.62.43.6-1.9-1.9-1.6-0.1
Social response proportion1.72.03.4-1.8-1.5-0.71.3
Social responses0.20.31.1-0.3-0.10.11.1
Social time0.40.51.3-0.2-0.10.91.1
Soleus weight1.44.25.4-1.9-1.9-2.3-2.1
Bone: stiffness4.26.99.0-3.0-2.8-2.7-1.7
Tibialis anterior weight0.10.30.6-0.3-0.2-0.8-0.6
Time to tail flick, vehicle, before self-admin0.20.20.3-0.4-0.5-0.3-0.4
Time to tail flick, vehicle, after self-admin0.50.62.0-0.3-0.1-0.6-1.4
Time to tail flick, test, before self-admin1.41.52.7-1.6-1.6-0.90.0
Time to tail flick, test, after self-admin0.60.60.80.90.40.90.9
Delta time to tail flick, vehicle, before/after SA0.20.20.40.60.60.1-0.4
Delta time to tail flick, test, before/after SA0.90.91.81.31.30.60.2
Tibia length0.51.31.8-1.3-1.4-1.2-0.2
Ambulatory time before self-admin1.01.41.7-1.0-1.2-1.2-1.3
Ambulatory time after self-admin1.72.02.6-1.6-1.6-1.3-1.0
Bone: tissue strength1.11.13.5-0.4-0.2-0.8-1.9
Total heroin consumption0.40.51.3-0.4-0.5-0.3-1.1
Bone: trabecular number0.40.61.5-0.8-0.6-1.20.1
Bone: trabecular spacing0.10.10.40.20.00.3-0.6
Bone: trabecular thickness0.40.51.1-0.5-0.3-0.71.0
Bone: trabecular tissue density0.40.61.20.91.10.3-0.5
Water consumed over 24 hours0.10.10.10.10.3-0.40.3
Number of responses in last shaping day0.30.30.4-0.6-0.6-0.60.5