Hey1

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 PL gene expression Hey1 0.22 0.12 1.6e-14 0.09 0.09 0.09 0.09 6.4e-10 5.9e-10 2.1e-10 4.3e-10
2 Adipose gene expression Hey1 0.04 0.03 1.6e-03 0 0 -0 -0 1.1e-01 1.7e-01 7.5e-01 4.5e-01
3 BLA gene expression Hey1 0.08 0.06 1.6e-03 0.02 0.03 0.01 0.03 4.5e-02 1.4e-02 8.9e-02 1.5e-02
4 Liver gene expression Hey1 0.03 0.03 6.4e-03 0 0.01 0.01 0 2.1e-01 1.8e-02 3.4e-02 7.2e-02

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 density0.91.62.01.2-1.2-1.21.4
Area under the delay curve0.90.91.6-1.00.50.9-1.3
BMI with tail3.712.116.4-4.04.04.0-0.4
BMI without tail8.317.223.1-4.84.64.8-1.5
Body weight1.66.59.5-2.83.13.0-0.0
Weight adjusted by age0.10.10.10.30.2-0.20.2
Bone surface0.81.43.90.7-0.8-0.72.0
Bone volume1.11.94.51.0-1.1-1.12.1
Breakpoint value in progressive ratio session0.30.40.6-0.70.80.6-0.0
Number of licking bursts0.10.10.1-0.30.00.3-0.3
Context. condit. distance diff. score0.20.20.7-0.30.30.3-0.8
Locomotion velocity, session 10.30.40.6-0.70.80.6-0.3
Locomotion distance, session 10.40.50.7-0.60.90.70.6
Locomotion velocity, session 21.41.72.3-1.51.51.5-0.1
Locomotion distance, session 20.81.12.4-0.60.90.81.6
Locomotion velocity, session 30.10.20.40.3-0.0-0.5-0.6
Locomotion distance, session 30.30.30.80.3-0.2-0.30.9
Stereotopy head waving bouts, day 30.10.20.20.4-0.4-0.50.3
Stereotopy head waving duration, day 30.40.40.7-0.50.60.5-0.8
Locomotion velocity, session 78.39.116.22.8-2.4-2.64.0
Locomotion distance, session 74.45.66.42.5-2.4-2.32.2
Stereotopy head waving bouts, day 70.60.71.0-1.01.01.0-0.1
Stereotopy head waving duration, day 71.21.41.7-1.31.31.2-0.9
Locomotion velocity, session 82.02.43.3-1.81.71.8-0.4
Locomotion distance, session 80.50.70.9-0.90.91.00.3
Degree of sensitization distance0.80.81.01.0-1.0-0.90.8
Degree of sensitization stereotypy2.22.43.2-1.81.71.7-1.0
Conditioned locomotion0.20.30.90.3-0.2-0.30.9
Cocaine response after cond. corrected1.31.52.21.4-1.5-1.30.2
Cocaine response after cond. not corrected3.23.54.72.2-2.1-2.10.5
Cocaine response before conditioning0.70.81.31.0-0.7-1.10.8
Saline control response0.60.82.50.5-0.4-0.61.6
Bone: connectivity density1.11.72.01.3-1.3-1.21.4
Bone: cortical apparent density0.20.30.4-0.50.70.5-0.4
Bone: cortical area1.01.52.2-1.51.31.5-0.0
Bone: cortical porosity0.10.10.1-0.20.30.30.1
Bone: cortical porosity0.00.00.0-0.10.20.10.2
Bone: cortical thickness0.71.21.6-1.31.21.3-0.2
Bone: cortical thickness4.24.96.6-2.62.42.4-1.3
Bone: cortical tissue density0.20.30.4-0.50.60.5-0.4
Condit. Reinf. active minus inactive responses2.93.24.8-1.51.91.42.2
Condit. Reinf. active-inactive response ratio3.94.37.0-1.72.21.62.6
Condit. Reinf. active responses2.63.14.7-1.51.91.42.2
Condit. Reinf. inactive responses0.30.40.5-0.70.70.7-0.2
Incentive salience index mean2.93.54.4-1.72.01.72.1
Condit. Reinf. lever presses1.62.22.9-1.51.61.71.1
Condit. Reinf. active minus inactive responses1.41.84.51.0-0.9-0.82.1
Condit. Reinf. active-inactive response ratio0.70.72.80.10.00.11.7
Condit. Reinf. active responses2.94.17.41.8-1.7-1.62.7
Condit. Reinf. inactive responses1.21.51.71.2-1.2-1.21.3
Condit. Reinf. lever presses0.81.01.71.0-0.9-0.81.3
Condit. Reinf. lever reinforcers received2.73.86.61.8-1.7-1.62.6
Conditioned reinforcement - actives2.42.43.11.8-1.3-1.61.5
Conditioned reinforcement - inactives1.11.32.21.5-0.8-1.20.9
Conditioned locomotion0.50.61.10.6-1.1-0.80.6
Patch foraging indifference point 0 sec0.81.21.71.3-1.2-1.3-0.1
Patch foraging indifference point AUC0.10.10.20.40.1-0.50.1
Delay disc. indifference point, 0s delay0.10.10.5-0.0-0.20.1-0.7
Delay disc. indifference point, 16s delay0.50.60.8-0.90.30.9-0.7
Delay disc. indifference point, 2s delay1.21.33.3-0.91.81.00.6
Delay disc. indifference point, 24s delay0.30.30.5-0.1-0.70.0-0.7
Delay disc. indifference point, 4s delay3.03.23.9-2.01.61.8-1.7
Delay disc. indifference point, 8s delay0.50.51.90.1-0.4-0.1-1.4
Indifference point function ln k0.91.21.9-1.10.61.2-1.4
Indifference point function log k0.91.21.9-1.10.61.2-1.4
Patch foraging total patch changes 0 sec0.50.91.21.1-1.0-1.1-0.5
Patch foraging total patch changes 12 sec0.91.11.91.4-0.7-1.3-0.7
Patch foraging total patch changes 18 sec0.20.20.6-0.10.8-0.1-0.4
Patch foraging total patch changes 24 sec0.20.20.40.60.0-0.6-0.4
Patch foraging total patch changes 6 sec0.10.20.40.5-0.2-0.6-0.4
Patch foraging time to switch 0 sec1.52.33.7-1.51.91.51.2
Patch foraging water rate 0 sec0.40.61.00.5-1.0-0.5-0.9
Patch foraging water rate 12 sec0.50.71.80.8-1.3-0.6-0.0
Patch foraging water rate 18 sec0.10.20.30.2-0.5-0.20.6
Patch foraging water rate 24 sec0.10.20.80.0-0.1-0.0-0.9
Patch foraging water rate 6 sec0.50.71.40.7-1.2-0.8-0.4
Seeking ratio, delayed vs. immediate footshock4.04.66.82.5-2.1-2.60.7
Distance traveled before self-admin1.81.92.2-1.51.21.5-1.3
Distance traveled after self-admin1.11.21.8-1.00.81.1-1.3
Extensor digitorum longus weight10.340.554.4-7.47.37.2-1.7
Bone: elastic displacement0.30.31.20.2-0.1-0.21.1
Bone: elastic work0.10.20.6-0.10.20.20.8
Bone: endosteal estimation0.71.53.1-1.00.91.01.7
Bone: endosteal perimeter0.10.20.9-0.1-0.10.20.9
Epididymis fat weight0.20.50.90.6-0.8-0.5-0.9
Delta time in closed arm before/after self-admin1.51.52.0-1.31.41.0-1.1
Delta time in open arm before/after self-admin0.30.30.50.7-0.6-0.60.3
Diff in mean of infusions in LGA sessions1.51.72.6-1.61.31.60.1
Delay discount exponential model param0.60.61.40.8-0.3-0.71.2
Extinction: sum of active levers before priming0.10.10.1-0.40.20.1-0.3
Active lever presses in extinction session 60.00.00.10.2-0.1-0.20.2
Bone: final force1.11.82.5-1.51.61.60.3
Bone: final moment0.91.52.0-1.41.41.40.3
Food consumed during 24 hour testing period1.72.02.6-1.61.51.4-1.1
Glucose1.42.23.5-1.51.91.3-1.1
Heart weight0.70.91.81.1-0.8-1.3-0.4
Delay discount hyperbolic model param0.90.91.51.0-0.6-1.01.2
Intermittent access intake day 1-15 change1.71.72.81.7-1.2-1.50.0
Intermittent access intake escalation1.21.11.91.4-1.1-1.20.1
Intermittent access intake escalation 23.03.04.92.2-1.7-2.00.6
Intermitt. access day 1 inactive lever presses0.40.41.6-0.20.10.1-1.3
Intermitt. access day 15 inactive lever presses2.83.14.12.0-1.8-2.0-1.1
Intermitt. access escalation Index0.20.20.7-0.30.00.1-0.9
Intermittent access day 1 total infusions4.14.76.12.4-2.5-2.01.7
Intermittent access day 15 total infusions6.57.310.03.2-3.0-2.71.5
Intermittent access terminal intake (last 3 days)6.77.310.13.2-3.1-2.81.2
Intermittent access total infusions5.66.39.22.9-3.0-2.51.3
Intermittent access day 1 locomotion0.80.91.41.1-1.2-0.80.5
Intermittent access day 15 total locomotion0.60.60.90.9-0.5-0.80.9
Intermittent access total locomotion1.61.82.81.6-1.7-1.30.7
Intermittent access standard deviation2.62.84.31.9-2.1-1.7-0.3
Left kidney weight0.51.32.30.8-1.3-0.8-1.5
Right kidney weight0.72.04.30.9-1.4-0.9-2.1
Cocaine induced anxiety0.10.20.50.2-0.1-0.3-0.7
Post-drug Anxiety0.50.50.8-0.90.80.5-0.5
Baseline Anxiety0.10.10.4-0.10.3-0.10.7
Tail length0.51.52.2-1.21.41.5-0.7
Length with tail0.00.10.10.30.0-0.10.4
Length without tail1.63.64.32.1-1.8-2.01.7
Times rat made contact with spout0.20.20.5-0.40.20.3-0.7
Lifetime Intake6.27.110.23.1-3.2-2.61.3
Liver weight, left1.62.53.9-1.82.01.7-0.3
Liver weight, right0.20.30.90.3-0.2-0.2-1.0
Locomotor activity1.22.02.8-1.31.51.7-1.0
Locomotor testing distance0.40.71.0-0.81.01.0-0.7
Locomotor testing rearing0.40.70.80.8-0.7-0.8-0.9
Locomotion in novel chamber1.82.23.2-1.61.81.70.2
Locomotion in novel chamber post-restriction0.20.20.8-0.10.20.0-0.9
Fecal boli incidents, locomotor time 10.20.20.40.4-0.5-0.40.7
Change in fecal boli incidents, locomotor task0.30.30.50.7-0.1-0.6-0.3
Time >=10cm from walls, locomotor time 10.00.00.10.20.3-0.20.2
Bouts of movement, locomotor time 11.41.64.80.4-1.0-0.6-2.2
Total resting periods, locomotor time 10.91.01.90.6-1.0-0.8-1.4
Rest time, locomotor task time 10.50.61.40.40.8-0.5-1.2
Distance moved, locomotor task time 10.40.40.7-0.8-0.10.80.4
Vertical activity count, locomotor time 11.31.42.2-1.50.91.50.5
Time >=10cm from walls, locomotor time 20.20.20.4-0.40.60.30.5
Bouts of movement, locomotor time 21.71.87.1-0.40.20.1-2.7
Total resting periods, locomotor time 22.22.39.0-0.40.50.1-3.0
Rest time, locomotor task time 21.21.33.5-0.41.30.2-1.9
Distance moved, locomotor task time 20.20.20.3-0.5-0.40.60.4
Vertical activity count, locomotor time 22.42.64.5-2.11.12.10.6
Light reinforcement 10.10.20.4-0.10.50.1-0.6
Light reinforcement 21.72.33.8-2.01.41.8-0.4
Bone: marrow area0.81.62.9-1.11.01.21.7
Bone: maximum diameter0.81.52.3-1.51.11.5-0.5
Bone: maximum force0.91.52.1-1.41.41.4-0.1
Bone: maximum moment0.71.11.6-1.31.21.2-0.1
Mean time between licks in bursts0.60.93.30.4-0.3-0.31.8
Mean num. licks in bursts0.00.00.00.00.2-0.1-0.1
Cd content in liver0.80.92.1-0.70.50.8-1.5
Co content in liver1.31.55.2-0.60.40.7-2.3
Cu content in liver2.53.15.21.9-2.3-1.80.0
Fe content in liver0.70.83.20.1-0.2-0.11.8
K content in liver1.21.21.81.3-1.3-1.10.4
Mg content in liver1.11.11.41.0-1.1-0.81.2
Mn content in liver0.20.20.50.4-0.3-0.40.7
Na content in liver0.10.10.4-0.00.0-0.0-0.6
Rb content in liver0.00.00.00.10.0-0.0-0.1
Se content in liver0.20.30.50.5-0.7-0.6-0.2
Sr content in liver0.10.10.1-0.20.10.10.4
Zn content in liver1.61.72.51.4-1.6-1.5-0.1
Bone: minimum diameter2.03.74.6-1.81.71.92.1
Time in familiar zone, hab. session 10.70.72.7-0.1-0.0-0.0-1.7
Time in novel zone, hab. session 10.70.70.80.9-0.7-0.90.8
Total zone transitions, hab. session 10.20.30.9-0.30.60.30.9
Total locomotion distance, hab. session 10.50.62.5-0.00.30.11.6
Locomotion velocity, hab. session 10.70.92.3-0.50.80.71.5
Time in familiar zone, hab. session 21.81.82.01.3-1.4-1.4-1.3
Time in novel zone, hab. session 20.90.91.2-0.91.10.81.0
Total zone transitions, hab. session 20.00.00.0-0.00.2-0.0-0.2
Total locomotion distance, hab. session 20.20.20.6-0.30.40.20.8
Locomotion velocity, hab. session 20.10.20.5-0.10.30.20.7
Novel to familiar place preference ratio0.70.71.1-1.01.00.9-0.2
Novelty place preference0.60.60.8-0.90.90.8-0.4
Time in novel zone, NPP test0.90.91.2-1.01.10.9-0.8
Total zone transitions, NPP test0.00.10.2-0.20.40.20.1
Total locomotion distance, NPP test0.20.20.4-0.30.50.30.6
Locomotion velocity, NPP test0.20.31.00.10.1-0.11.0
Intraocular pressure2.33.15.0-2.22.01.9-0.4
Delta distance traveled before/after self-admin0.20.20.30.5-0.5-0.40.5
Delta ambulatory time before/after self-admin0.91.01.51.1-1.2-1.00.5
Parametrial fat weight0.10.10.2-0.30.30.2-0.4
Pavlov. Cond. lever latency0.70.91.30.7-0.9-1.1-1.0
Pavlov. Cond. magazine entry latency0.50.60.9-0.81.00.80.5
Pavlov. Cond. change in total contacts2.93.14.5-1.92.02.10.4
Pavlov. Cond. index score0.91.11.7-1.01.21.30.4
Pavlov. Cond. latency score0.70.91.6-0.91.11.30.4
Pavlov. Cond. lever contacts1.11.32.1-0.81.01.21.5
Pavlov. Cond. magazine entry number1.21.42.11.4-1.5-1.30.1
Pavlov. Cond. intertrial magazine entries2.42.93.71.9-1.9-1.81.0
Pavlov. Cond. lever-magazine prob. diff.0.91.21.9-1.11.21.40.2
Pavlov. Cond. response bias1.41.72.4-1.31.61.40.7
Pavlov. Cond. lever latency0.81.11.5-1.21.21.1-0.6
Pavlov. Cond. magazine entry latency0.10.10.1-0.40.30.40.2
Pavlov. Cond. change in total contacts0.40.62.30.0-0.10.11.5
Pavlov. Cond. index score0.20.20.90.1-0.1-0.10.9
Pavlov. Cond. latency score0.10.20.50.2-0.3-0.20.7
Pavlov. Cond. lever contacts0.71.01.21.0-1.1-1.00.9
Pavlov. Cond. magazine entry number0.30.30.40.6-0.7-0.6-0.4
Pavlov. Cond. intertrial magazine entries0.10.20.2-0.40.40.50.4
Pavlov. Cond. lever-magazine prob. diff.0.20.30.80.3-0.3-0.30.9
Pavlov. Cond. response bias0.40.51.60.4-0.4-0.31.2
Time in closed arm before self-admin1.92.23.11.8-1.6-1.60.7
Time in closed arm after self-admin0.30.31.0-0.30.10.3-1.0
Time in open arm before self-admin0.80.91.2-1.11.11.00.4
Time in open arm after self-admin0.70.81.6-0.91.30.90.1
Bone: periosteal estimation1.42.93.5-1.81.71.91.3
Bone: periosteal perimeter0.30.50.9-0.90.71.00.2
Bone: post-yield work0.71.01.31.0-1.1-1.1-0.8
Progressive ratio test 1 active lever presses4.04.26.32.4-2.5-2.1-0.4
Progressive ratio test 1 breakpoint3.73.96.02.4-2.5-2.0-0.2
Progressive ratio test 1 inactive lever presses0.60.71.10.9-1.0-0.8-0.5
Progressive ratio test 2 active lever presses2.32.54.02.0-1.5-1.70.9
Progressive ratio test 2 breakpoint2.02.33.61.9-1.4-1.61.1
Progressive ratio test 2 inactive lever presses1.61.82.91.6-1.7-1.40.1
Incentive Sensitization - Responses1.71.73.61.3-0.7-1.11.9
Incentive Sensitization - Breakpoint1.51.52.91.3-0.7-1.11.7
Sum of active levers in priming session hrs 5-60.40.40.70.5-0.4-0.60.8
Progressive ratio1.01.13.5-0.30.80.4-1.9
Food seeking constrained by brief footshock3.34.15.3-2.32.12.31.1
Seeking ratio, punishment vs. effort1.51.86.4-0.60.20.62.5
Retroperitoneal fat weight0.10.30.40.6-0.5-0.60.2
Reaction time number correct0.30.41.00.5-0.4-0.41.0
Reaction time mean minus median0.81.12.8-0.60.80.7-1.7
Reaction time mean minus median AUC1.21.62.2-1.01.41.0-1.5
Reaction time num false alarms0.81.13.50.50.7-0.51.9
Reaction time num false alarms AUC0.91.23.10.70.8-0.71.8
Reaction time trials correct on left0.30.41.00.5-0.4-0.41.0
Reaction time trials on left0.30.41.10.6-0.4-0.41.0
Reaction time mean1.01.41.9-1.31.41.3-0.3
Reaction time mean AUC1.11.52.0-1.41.41.4-0.3
Median of all reaction times1.21.72.3-1.51.51.50.2
Reaction time omissions0.20.20.3-0.50.50.6-0.2
Reaction time false alarm rate0.30.41.20.11.1-0.20.6
Reaction time premature initiation rate0.91.21.81.1-1.3-1.0-0.9
Reaction time premature initiations0.91.32.11.3-1.4-1.2-0.4
Std. dev. reaction times0.81.12.4-0.80.80.9-1.5
Reaction time trials completed0.30.41.10.6-0.4-0.41.0
Reaction time trials AUC0.40.51.40.6-0.4-0.41.2
Run reversals in cocaine runway, females3.13.04.7-2.01.62.2-1.0
Run reversals in cocaine runway, males3.43.35.7-1.92.42.00.1
Latency to leave start box in cocaine runway0.60.61.40.6-1.2-0.70.2
Latency to leave start box in cocaine runway, F2.62.84.81.7-2.2-1.80.5
Latency to leave start box in cocaine runway, M0.50.50.7-0.80.60.80.1
Std. dev. time between licks in bursts0.50.51.70.4-0.1-0.51.3
Total sessions with >9 infusions3.03.35.12.3-2.2-1.80.3
Short access day 1 total inactive lever presses1.71.93.61.9-1.3-1.60.0
Short access day 10 total inactive lever presses0.50.62.2-0.4-0.10.0-1.5
Short access day 1 total infusions2.83.04.72.0-2.2-1.70.1
Short access day 10 total infusions5.76.310.52.8-3.2-2.60.4
Short access total infusions6.37.111.03.0-3.3-2.71.0
Short access day 1 locomotion0.50.51.50.4-0.6-0.2-1.2
Short access day 10 total locomotion0.10.10.1-0.2-0.20.3-0.3
Short access total locomotion0.70.71.70.8-1.3-0.6-0.4
One hour access (0.3 mA shock)2.52.94.72.2-1.9-1.61.0
Compulsive drug intake3.13.25.2-2.21.62.3-0.3
One hour access (shock baseline)6.17.110.73.3-2.9-2.90.9
Social response proportion0.40.41.2-0.10.5-0.0-1.1
Social responses0.91.23.0-0.51.20.2-1.7
Social time0.91.22.51.4-0.5-1.60.4
Soleus weight1.44.06.0-2.42.02.2-1.0
Bone: stiffness0.30.40.6-0.80.60.6-0.5
Tibialis anterior weight9.934.045.6-6.76.86.7-0.1
Time to tail flick, vehicle, before self-admin0.70.70.70.8-0.8-0.90.8
Time to tail flick, vehicle, after self-admin1.41.51.91.2-1.3-1.11.4
Time to tail flick, test, before self-admin0.20.20.30.3-0.5-0.2-0.5
Time to tail flick, test, after self-admin1.71.72.61.5-1.6-1.4-0.1
Delta time to tail flick, vehicle, before/after SA1.21.21.41.2-1.1-1.01.1
Delta time to tail flick, test, before/after SA0.30.30.60.4-0.4-0.4-0.8
Tibia length0.51.52.21.3-1.5-1.4-0.3
Ambulatory time before self-admin1.21.21.5-1.11.21.1-1.0
Ambulatory time after self-admin0.10.10.2-0.30.20.4-0.4
Bone: tissue strength1.31.31.91.4-1.2-1.4-0.1
Total heroin consumption0.10.10.4-0.00.2-0.10.6
Bone: trabecular number0.40.60.80.7-0.9-0.80.7
Bone: trabecular spacing0.20.20.3-0.60.50.5-0.1
Bone: trabecular thickness1.01.21.51.0-1.2-1.11.0
Bone: trabecular tissue density0.40.51.8-0.30.3-0.1-1.4
Water consumed over 24 hours0.50.70.9-1.00.90.8-0.6
Number of responses in last shaping day0.20.20.70.1-0.10.1-0.8