Gulp1

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 Brain mRNA stability Gulp1 0.06 0.05 6.5e-06 0.06 0.05 0.06 0.06 5.7e-06 1.6e-05 2.2e-06 4.8e-06
2 Brain intron excision ratio chr9_46884963_46897134 0.03 0.03 8.8e-03 0.01 0.01 -0 0 7.7e-02 3.7e-02 3.7e-01 2.8e-01
3 Adipose gene expression Gulp1 0.05 0.04 6.9e-06 0.03 0.03 0.03 0.03 2.4e-04 4.1e-04 6.0e-04 5.5e-04
4 NAcc gene expression Gulp1 0.4 0.15 0.0e+00 0.27 0.27 0.28 0.28 3.0e-31 8.1e-32 4.9e-32 3.0e-32

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 density3.86.410.2-3.22.3-2.71.6
Area under the delay curve0.80.92.80.60.20.5-1.7
BMI with tail2.88.913.53.7-3.33.4-0.4
BMI without tail0.00.00.1-0.20.2-0.2-0.0
Body weight0.10.50.50.7-0.60.7-0.7
Weight adjusted by age1.82.03.70.7-1.90.71.8
Bone surface0.50.92.8-0.7-0.2-0.51.7
Bone volume1.52.63.9-2.01.0-1.61.7
Breakpoint value in progressive ratio session0.30.30.6-0.60.8-0.5-0.2
Number of licking bursts0.30.40.7-0.80.6-0.70.3
Context. condit. distance diff. score1.31.62.7-1.41.6-1.1-0.8
Locomotion velocity, session 10.60.72.30.20.60.2-1.5
Locomotion distance, session 11.21.52.21.5-0.41.2-1.5
Locomotion velocity, session 23.84.46.72.6-2.42.30.1
Locomotion distance, session 23.65.18.52.9-2.22.6-0.4
Locomotion velocity, session 30.10.20.3-0.40.5-0.50.0
Locomotion distance, session 30.10.10.2-0.40.3-0.40.1
Stereotopy head waving bouts, day 30.30.40.5-0.70.7-0.6-0.0
Stereotopy head waving duration, day 30.40.40.8-0.40.9-0.3-0.8
Locomotion velocity, session 70.70.81.20.7-1.10.70.9
Locomotion distance, session 70.91.11.61.2-1.21.10.2
Stereotopy head waving bouts, day 70.30.30.5-0.50.7-0.4-0.5
Stereotopy head waving duration, day 70.50.60.9-0.71.0-0.7-0.7
Locomotion velocity, session 80.10.10.10.2-0.30.20.4
Locomotion distance, session 80.71.01.51.2-0.91.2-0.3
Degree of sensitization distance1.41.42.11.5-1.21.4-0.5
Degree of sensitization stereotypy0.10.10.1-0.30.3-0.4-0.1
Conditioned locomotion0.40.51.6-0.20.7-0.1-1.3
Cocaine response after cond. corrected0.70.81.21.0-0.50.8-1.1
Cocaine response after cond. not corrected0.70.71.30.9-0.40.8-1.2
Cocaine response before conditioning0.00.00.0-0.00.1-0.00.0
Saline control response0.10.20.20.4-0.30.4-0.5
Bone: connectivity density0.91.42.0-1.30.6-1.11.4
Bone: cortical apparent density2.12.84.61.8-2.11.70.5
Bone: cortical area0.30.40.60.6-0.80.70.2
Bone: cortical porosity1.31.42.3-1.51.1-1.40.5
Bone: cortical porosity1.61.82.7-1.71.4-1.50.3
Bone: cortical thickness0.30.50.70.7-0.80.8-0.1
Bone: cortical thickness0.10.20.30.4-0.40.5-0.1
Bone: cortical tissue density2.02.74.41.8-2.11.60.6
Condit. Reinf. active minus inactive responses2.32.58.20.41.30.5-2.9
Condit. Reinf. active-inactive response ratio3.13.310.50.01.70.2-3.2
Condit. Reinf. active responses1.21.44.50.41.00.4-2.1
Condit. Reinf. inactive responses0.40.40.90.9-0.60.7-0.3
Incentive salience index mean1.01.23.40.01.20.1-1.8
Condit. Reinf. lever presses0.50.71.5-0.81.2-0.6-0.5
Condit. Reinf. active minus inactive responses0.91.12.0-0.71.4-0.6-1.3
Condit. Reinf. active-inactive response ratio1.41.52.7-1.21.7-1.1-0.7
Condit. Reinf. active responses0.30.41.10.40.20.5-1.1
Condit. Reinf. inactive responses0.91.11.71.1-1.31.20.1
Condit. Reinf. lever presses0.20.30.5-0.70.7-0.5-0.1
Condit. Reinf. lever reinforcers received0.40.51.40.60.10.6-1.2
Conditioned reinforcement - actives0.20.20.5-0.30.70.2-0.7
Conditioned reinforcement - inactives0.60.71.1-0.71.1-0.4-0.9
Conditioned locomotion1.31.52.71.3-1.31.6-0.1
Patch foraging indifference point 0 sec1.82.87.60.2-1.90.22.7
Patch foraging indifference point AUC1.11.32.21.5-1.21.2-0.1
Delay disc. indifference point, 0s delay0.10.10.40.3-0.70.00.3
Delay disc. indifference point, 16s delay0.70.72.0-0.10.90.1-1.4
Delay disc. indifference point, 2s delay0.10.10.30.4-0.50.1-0.2
Delay disc. indifference point, 24s delay0.90.91.3-0.61.1-0.6-1.2
Delay disc. indifference point, 4s delay0.70.71.50.9-0.30.8-1.2
Delay disc. indifference point, 8s delay0.20.20.80.20.20.1-0.9
Indifference point function ln k0.20.20.70.3-0.80.40.3
Indifference point function log k0.20.20.70.3-0.80.40.3
Patch foraging total patch changes 0 sec2.03.37.30.6-2.30.52.7
Patch foraging total patch changes 12 sec0.81.03.20.2-0.90.11.8
Patch foraging total patch changes 18 sec2.12.74.91.5-2.21.31.4
Patch foraging total patch changes 24 sec0.81.02.40.3-1.20.21.5
Patch foraging total patch changes 6 sec1.31.95.9-0.0-1.3-0.02.4
Patch foraging time to switch 0 sec0.50.81.2-1.01.1-0.90.2
Patch foraging water rate 0 sec0.50.81.31.0-1.10.9-0.4
Patch foraging water rate 12 sec0.71.02.70.0-1.20.01.6
Patch foraging water rate 18 sec0.60.81.2-1.10.5-1.00.9
Patch foraging water rate 24 sec0.00.00.0-0.10.10.1-0.2
Patch foraging water rate 6 sec0.30.41.2-0.0-0.80.11.1
Seeking ratio, delayed vs. immediate footshock0.10.10.3-0.20.6-0.3-0.4
Distance traveled before self-admin0.10.10.5-0.2-0.0-0.10.7
Distance traveled after self-admin2.32.63.8-1.91.0-1.71.6
Extensor digitorum longus weight5.220.532.25.7-5.04.9-0.9
Bone: elastic displacement0.50.61.4-0.80.2-0.61.2
Bone: elastic work0.20.30.5-0.70.6-0.50.3
Bone: endosteal estimation0.00.00.0-0.10.1-0.10.2
Bone: endosteal perimeter0.00.00.00.10.10.2-0.2
Epididymis fat weight0.00.10.3-0.0-0.2-0.10.6
Delta time in closed arm before/after self-admin1.61.54.2-1.0-0.0-1.02.0
Delta time in open arm before/after self-admin0.20.20.2-0.20.5-0.4-0.5
Diff in mean of infusions in LGA sessions1.11.21.8-1.31.0-1.30.5
Delay discount exponential model param0.90.93.2-0.5-0.4-0.41.8
Extinction: sum of active levers before priming0.10.10.30.20.20.2-0.5
Active lever presses in extinction session 61.61.53.01.4-0.41.0-1.7
Bone: final force0.10.20.30.4-0.30.4-0.5
Bone: final moment0.10.20.20.5-0.40.5-0.4
Food consumed during 24 hour testing period0.60.71.11.0-0.70.9-0.6
Glucose0.10.10.2-0.10.4-0.2-0.4
Heart weight0.40.50.90.7-0.90.70.5
Delay discount hyperbolic model param0.91.03.0-0.7-0.1-0.61.7
Intermittent access intake day 1-15 change0.50.50.8-0.80.9-0.7-0.2
Intermittent access intake escalation1.41.42.3-1.21.5-1.1-0.8
Intermittent access intake escalation 22.02.03.8-1.41.9-1.2-0.9
Intermitt. access day 1 inactive lever presses0.00.00.1-0.20.3-0.20.0
Intermitt. access day 15 inactive lever presses0.30.30.5-0.60.5-0.20.7
Intermitt. access escalation Index1.00.92.4-0.31.1-0.1-1.6
Intermittent access day 1 total infusions0.20.20.50.20.20.5-0.7
Intermittent access day 15 total infusions1.11.22.3-1.11.5-0.7-1.0
Intermittent access terminal intake (last 3 days)0.40.40.8-0.50.90.0-0.8
Intermittent access total infusions0.30.40.7-0.50.8-0.1-0.7
Intermittent access day 1 locomotion0.10.10.40.2-0.20.6-0.0
Intermittent access day 15 total locomotion0.10.10.20.00.20.2-0.4
Intermittent access total locomotion0.10.10.40.3-0.10.6-0.3
Intermittent access standard deviation0.50.51.0-0.91.0-0.5-0.3
Left kidney weight0.20.51.5-0.5-0.3-0.41.2
Right kidney weight0.20.61.8-0.3-0.5-0.21.3
Cocaine induced anxiety0.91.01.61.3-1.21.1-0.2
Post-drug Anxiety2.12.33.41.8-1.01.6-1.5
Baseline Anxiety0.60.72.10.00.80.1-1.4
Tail length1.75.07.3-2.52.7-2.3-1.0
Length with tail0.51.82.6-1.61.5-1.4-0.6
Length without tail1.02.23.01.7-1.71.6-0.1
Times rat made contact with spout0.60.71.2-0.81.1-0.7-0.5
Lifetime Intake0.20.20.5-0.10.40.3-0.7
Liver weight, left1.21.96.2-0.7-0.6-0.72.5
Liver weight, right0.20.40.6-0.80.3-0.70.6
Locomotor activity0.10.20.2-0.50.3-0.40.4
Locomotor testing distance1.11.82.6-1.61.0-1.41.2
Locomotor testing rearing1.82.84.2-2.12.0-1.70.1
Locomotion in novel chamber5.97.211.23.4-2.53.1-1.4
Locomotion in novel chamber post-restriction4.04.87.42.7-2.12.6-0.8
Fecal boli incidents, locomotor time 12.42.64.52.1-1.41.8-1.0
Change in fecal boli incidents, locomotor task0.40.40.7-0.60.2-0.70.8
Time >=10cm from walls, locomotor time 11.91.93.21.8-1.61.40.1
Bouts of movement, locomotor time 18.18.813.33.6-2.13.4-2.4
Total resting periods, locomotor time 18.18.913.93.7-2.33.4-2.2
Rest time, locomotor task time 11.41.52.21.3-0.31.5-1.4
Distance moved, locomotor task time 10.70.81.9-0.8-0.1-0.91.4
Vertical activity count, locomotor time 11.41.52.3-1.50.7-1.31.2
Time >=10cm from walls, locomotor time 20.10.10.20.1-0.30.20.5
Bouts of movement, locomotor time 23.03.35.42.2-1.52.3-0.8
Total resting periods, locomotor time 22.82.94.92.1-1.62.2-0.4
Rest time, locomotor task time 20.00.00.1-0.30.3-0.10.0
Distance moved, locomotor task time 20.20.20.50.3-0.70.10.5
Vertical activity count, locomotor time 20.50.50.80.7-0.20.9-0.9
Light reinforcement 13.46.210.02.8-3.22.60.5
Light reinforcement 22.43.25.71.6-2.41.41.6
Bone: marrow area0.00.00.0-0.10.1-0.10.1
Bone: maximum diameter0.10.10.4-0.1-0.1-0.10.6
Bone: maximum force0.20.30.40.6-0.50.7-0.3
Bone: maximum moment0.20.30.50.7-0.60.7-0.2
Mean time between licks in bursts1.92.64.71.3-2.21.21.7
Mean num. licks in bursts0.30.40.70.8-0.50.7-0.4
Cd content in liver0.10.10.1-0.20.4-0.3-0.1
Co content in liver1.72.03.9-0.71.8-0.7-2.0
Cu content in liver1.61.93.2-0.91.8-0.8-1.8
Fe content in liver1.92.33.8-2.01.6-1.60.3
K content in liver1.91.93.41.2-1.81.11.3
Mg content in liver0.60.61.00.5-0.90.41.0
Mn content in liver0.20.30.50.5-0.70.50.3
Na content in liver2.02.26.5-1.1-0.3-0.92.6
Rb content in liver2.93.65.22.3-1.22.0-1.8
Se content in liver0.30.40.70.8-0.60.80.1
Sr content in liver0.00.00.00.10.0-0.0-0.1
Zn content in liver0.20.20.6-0.00.5-0.0-0.7
Bone: minimum diameter0.40.81.50.9-1.20.90.4
Time in familiar zone, hab. session 10.70.62.10.10.60.3-1.4
Time in novel zone, hab. session 11.21.21.7-1.30.7-1.31.0
Total zone transitions, hab. session 10.20.31.0-0.3-0.1-0.31.0
Total locomotion distance, hab. session 10.71.01.41.2-1.21.10.2
Locomotion velocity, hab. session 10.81.11.71.1-1.31.10.5
Time in familiar zone, hab. session 20.30.30.5-0.70.6-0.50.2
Time in novel zone, hab. session 20.10.10.30.00.1-0.1-0.5
Total zone transitions, hab. session 20.40.50.91.0-0.50.8-0.5
Total locomotion distance, hab. session 21.52.03.01.7-0.81.7-1.3
Locomotion velocity, hab. session 21.31.82.71.6-0.91.6-1.0
Novel to familiar place preference ratio0.10.10.40.2-0.00.2-0.6
Novelty place preference0.20.20.60.10.10.1-0.8
Time in novel zone, NPP test0.20.20.5-0.10.3-0.1-0.7
Total zone transitions, NPP test0.50.81.31.0-0.10.9-1.1
Total locomotion distance, NPP test0.71.01.61.3-0.81.3-0.4
Locomotion velocity, NPP test1.01.52.31.5-1.21.5-0.2
Intraocular pressure2.23.14.72.2-1.32.0-1.4
Delta distance traveled before/after self-admin0.70.71.0-1.00.3-1.01.0
Delta ambulatory time before/after self-admin0.70.71.5-0.90.1-0.81.2
Parametrial fat weight0.10.20.40.6-0.50.50.1
Pavlov. Cond. lever latency0.50.61.20.8-1.10.70.5
Pavlov. Cond. magazine entry latency0.20.20.5-0.40.7-0.3-0.4
Pavlov. Cond. change in total contacts0.30.30.80.60.20.4-0.9
Pavlov. Cond. index score0.30.40.8-0.50.9-0.4-0.4
Pavlov. Cond. latency score0.40.51.0-0.61.0-0.5-0.4
Pavlov. Cond. lever contacts0.20.20.5-0.50.7-0.3-0.3
Pavlov. Cond. magazine entry number0.81.01.91.1-1.41.00.3
Pavlov. Cond. intertrial magazine entries0.70.81.80.9-0.00.8-1.4
Pavlov. Cond. lever-magazine prob. diff.0.10.20.4-0.10.6-0.1-0.6
Pavlov. Cond. response bias0.20.20.5-0.40.7-0.4-0.4
Pavlov. Cond. lever latency0.60.81.51.2-0.81.0-0.5
Pavlov. Cond. magazine entry latency0.70.81.4-1.20.8-1.10.4
Pavlov. Cond. change in total contacts0.60.81.3-1.10.5-1.00.9
Pavlov. Cond. index score0.50.71.2-1.10.5-0.90.7
Pavlov. Cond. latency score0.40.50.9-0.90.4-0.80.7
Pavlov. Cond. lever contacts1.62.13.5-1.91.0-1.61.2
Pavlov. Cond. magazine entry number1.31.62.61.6-0.91.4-0.9
Pavlov. Cond. intertrial magazine entries3.74.87.42.7-1.92.6-1.3
Pavlov. Cond. lever-magazine prob. diff.0.40.50.8-0.90.3-0.80.7
Pavlov. Cond. response bias2.23.04.7-2.21.1-1.91.5
Time in closed arm before self-admin0.50.62.10.30.30.2-1.4
Time in closed arm after self-admin0.80.91.3-1.10.5-1.20.8
Time in open arm before self-admin0.40.40.60.6-0.70.80.5
Time in open arm after self-admin0.20.20.3-0.50.6-0.30.2
Bone: periosteal estimation0.00.00.0-0.0-0.10.00.2
Bone: periosteal perimeter0.00.00.10.2-0.10.3-0.3
Bone: post-yield work0.20.30.4-0.50.6-0.60.0
Progressive ratio test 1 active lever presses0.60.61.10.8-0.31.1-0.8
Progressive ratio test 1 breakpoint0.60.61.30.8-0.41.1-0.6
Progressive ratio test 1 inactive lever presses0.10.10.2-0.20.1-0.10.5
Progressive ratio test 2 active lever presses0.50.51.0-0.41.00.2-1.0
Progressive ratio test 2 breakpoint0.40.41.2-0.10.70.3-1.1
Progressive ratio test 2 inactive lever presses0.30.40.8-0.90.8-0.40.1
Incentive Sensitization - Responses0.80.81.7-0.91.3-0.4-0.6
Incentive Sensitization - Breakpoint0.60.61.4-0.81.2-0.3-0.6
Sum of active levers in priming session hrs 5-60.10.10.2-0.40.4-0.30.2
Progressive ratio2.73.15.21.9-0.61.8-2.3
Food seeking constrained by brief footshock0.00.00.00.1-0.10.1-0.1
Seeking ratio, punishment vs. effort1.41.73.8-1.30.2-1.21.9
Retroperitoneal fat weight0.31.01.51.1-1.21.10.3
Reaction time number correct0.81.11.81.2-1.41.1-0.0
Reaction time mean minus median1.11.52.4-1.61.3-1.30.6
Reaction time mean minus median AUC1.31.72.5-1.61.5-1.30.3
Reaction time num false alarms1.31.82.91.7-1.21.5-0.7
Reaction time num false alarms AUC1.92.64.12.0-1.71.8-0.4
Reaction time trials correct on left0.81.11.81.2-1.41.1-0.0
Reaction time trials on left0.71.01.61.1-1.31.1-0.1
Reaction time mean2.63.65.9-2.12.4-2.0-0.4
Reaction time mean AUC2.12.95.0-1.82.2-1.7-0.7
Median of all reaction times2.43.35.9-1.92.4-1.7-0.9
Reaction time omissions0.91.21.9-1.41.3-1.20.0
Reaction time false alarm rate0.50.71.31.1-0.81.0-0.3
Reaction time premature initiation rate0.10.10.1-0.40.2-0.40.2
Reaction time premature initiations0.10.10.20.3-0.40.30.2
Std. dev. reaction times2.23.14.8-2.21.8-1.90.7
Reaction time trials completed0.71.01.61.1-1.31.1-0.1
Reaction time trials AUC0.70.91.41.1-1.21.0-0.2
Run reversals in cocaine runway, females0.90.92.01.1-0.31.4-0.5
Run reversals in cocaine runway, males0.60.61.00.9-0.71.0-0.3
Latency to leave start box in cocaine runway2.52.74.22.0-1.71.9-0.3
Latency to leave start box in cocaine runway, F0.60.61.00.6-0.90.71.0
Latency to leave start box in cocaine runway, M3.63.55.52.3-1.72.0-1.4
Std. dev. time between licks in bursts1.01.22.20.9-1.50.91.0
Total sessions with >9 infusions1.71.93.21.6-1.31.8-0.5
Short access day 1 total inactive lever presses0.70.82.8-0.5-0.3-0.11.7
Short access day 10 total inactive lever presses0.70.71.1-0.81.0-1.1-0.4
Short access day 1 total infusions0.30.30.40.3-0.70.50.6
Short access day 10 total infusions0.40.51.40.10.50.5-1.2
Short access total infusions0.60.71.30.9-0.51.1-0.6
Short access day 1 locomotion0.10.10.40.2-0.60.20.3
Short access day 10 total locomotion0.20.20.5-0.40.00.00.7
Short access total locomotion0.40.40.80.7-0.80.90.0
One hour access (0.3 mA shock)1.31.62.9-0.81.7-0.5-1.6
Compulsive drug intake1.21.21.8-1.41.1-1.10.7
One hour access (shock baseline)1.11.33.6-0.31.1-0.3-1.9
Social response proportion0.80.91.3-0.71.1-0.6-1.1
Social responses1.31.62.9-1.21.7-0.9-1.2
Social time3.85.07.9-2.82.3-2.60.5
Soleus weight1.02.84.61.7-2.21.41.3
Bone: stiffness0.20.30.70.5-0.10.4-0.8
Tibialis anterior weight2.06.79.83.1-3.12.70.3
Time to tail flick, vehicle, before self-admin0.20.20.40.6-0.20.5-0.6
Time to tail flick, vehicle, after self-admin0.30.30.90.00.5-0.1-0.9
Time to tail flick, test, before self-admin0.60.61.01.0-0.60.7-0.7
Time to tail flick, test, after self-admin2.02.03.31.8-1.11.5-1.2
Delta time to tail flick, vehicle, before/after SA0.10.10.1-0.40.3-0.3-0.1
Delta time to tail flick, test, before/after SA0.10.10.20.4-0.30.3-0.1
Tibia length0.10.30.4-0.60.5-0.60.4
Ambulatory time before self-admin0.00.00.10.1-0.00.30.1
Ambulatory time after self-admin0.80.93.0-0.6-0.3-0.51.7
Bone: tissue strength0.20.20.50.4-0.00.2-0.7
Total heroin consumption0.10.10.40.10.20.1-0.7
Bone: trabecular number3.04.77.3-2.72.0-2.31.4
Bone: trabecular spacing2.43.24.82.2-1.32.0-1.4
Bone: trabecular thickness4.75.68.9-3.02.5-2.60.7
Bone: trabecular tissue density0.20.30.4-0.60.7-0.4-0.4
Water consumed over 24 hours0.40.51.40.60.10.5-1.2
Number of responses in last shaping day0.40.40.7-0.70.5-0.80.2