Kremen1

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 gene expression Kremen1 0.04 0.03 1.9e-04 0.04 0.03 0.04 0.04 1.8e-04 4.5e-04 9.7e-05 1.1e-04
2 Adipose gene expression Kremen1 0.13 0.09 3.2e-08 0.05 0.05 0.05 0.05 1.8e-06 2.6e-06 1.9e-06 1.8e-06
3 BLA gene expression Kremen1 0.16 0.09 3.1e-08 0.1 0.14 0.11 0.11 9.0e-06 1.1e-07 1.2e-06 1.7e-06
4 Liver gene expression Kremen1 0.04 0.03 5.0e-03 0.01 0 0 0 1.3e-02 7.3e-02 3.5e-01 9.5e-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 density0.10.20.4-0.50.60.4-0.2
Area under the delay curve4.04.25.52.3-2.2-2.0-1.6
BMI with tail0.00.00.0-0.20.10.10.1
BMI without tail0.10.10.5-0.00.20.3-0.7
Body weight1.45.76.82.6-2.3-2.3-2.3
Weight adjusted by age1.01.22.41.1-0.8-0.7-1.5
Bone surface0.20.30.40.5-0.3-0.6-0.6
Bone volume0.10.20.50.4-0.2-0.5-0.7
Breakpoint value in progressive ratio session1.71.82.31.5-1.4-1.3-1.1
Number of licking bursts3.13.64.32.1-1.8-1.9-1.8
Context. condit. distance diff. score4.14.96.2-2.52.32.31.8
Locomotion velocity, session 10.60.62.2-0.1-0.3-0.31.5
Locomotion distance, session 10.60.81.2-1.01.11.00.1
Locomotion velocity, session 20.50.62.1-0.3-0.1-0.31.4
Locomotion distance, session 22.13.03.8-2.01.71.41.8
Locomotion velocity, session 32.83.24.3-2.12.11.81.0
Locomotion distance, session 30.00.00.0-0.10.2-0.2-0.1
Stereotopy head waving bouts, day 31.01.21.5-1.21.21.10.6
Stereotopy head waving duration, day 31.82.12.5-1.61.41.51.3
Locomotion velocity, session 70.10.10.20.3-0.4-0.4-0.1
Locomotion distance, session 70.50.61.0-0.80.70.51.0
Stereotopy head waving bouts, day 70.30.30.4-0.60.60.70.4
Stereotopy head waving duration, day 70.10.10.2-0.00.20.2-0.5
Locomotion velocity, session 82.12.54.0-1.71.31.22.0
Locomotion distance, session 88.612.615.8-4.03.63.53.0
Degree of sensitization distance2.12.13.1-1.51.21.31.8
Degree of sensitization stereotypy0.20.20.20.5-0.4-0.3-0.5
Conditioned locomotion1.11.31.9-1.21.31.40.0
Cocaine response after cond. corrected2.12.43.01.7-1.6-1.4-1.4
Cocaine response after cond. not corrected2.72.93.61.9-1.6-1.4-1.9
Cocaine response before conditioning0.30.30.5-0.60.70.70.1
Saline control response1.01.31.7-1.31.10.91.3
Bone: connectivity density0.20.40.6-0.70.80.60.0
Bone: cortical apparent density0.81.01.21.1-1.0-1.0-1.0
Bone: cortical area3.24.85.92.4-2.2-2.1-2.1
Bone: cortical porosity0.20.20.50.3-0.2-0.1-0.7
Bone: cortical porosity0.10.10.2-0.30.40.4-0.0
Bone: cortical thickness2.23.65.42.0-1.6-1.6-2.3
Bone: cortical thickness0.50.60.9-0.71.00.9-0.5
Bone: cortical tissue density0.81.11.31.2-1.0-1.0-0.9
Condit. Reinf. active minus inactive responses0.60.71.10.6-0.9-1.10.6
Condit. Reinf. active-inactive response ratio1.01.14.2-0.3-0.3-0.42.1
Condit. Reinf. active responses0.20.20.50.3-0.5-0.70.4
Condit. Reinf. inactive responses0.30.41.30.4-0.1-0.3-1.1
Incentive salience index mean0.70.81.50.9-1.0-1.20.0
Condit. Reinf. lever presses0.10.20.20.4-0.3-0.4-0.5
Condit. Reinf. active minus inactive responses0.60.70.9-0.90.80.70.9
Condit. Reinf. active-inactive response ratio0.10.10.2-0.30.20.10.4
Condit. Reinf. active responses0.81.11.4-1.10.90.91.2
Condit. Reinf. inactive responses0.91.11.5-1.10.90.91.2
Condit. Reinf. lever presses0.60.81.0-1.01.00.90.6
Condit. Reinf. lever reinforcers received0.70.91.1-1.10.90.81.1
Conditioned reinforcement - actives0.00.00.00.1-0.2-0.00.1
Conditioned reinforcement - inactives0.10.10.20.3-0.4-0.3-0.0
Conditioned locomotion0.10.20.30.4-0.4-0.1-0.6
Patch foraging indifference point 0 sec2.03.23.71.9-1.8-1.7-1.7
Patch foraging indifference point AUC0.10.10.20.3-0.3-0.40.0
Delay disc. indifference point, 0s delay1.21.31.71.3-1.1-1.0-1.1
Delay disc. indifference point, 16s delay3.13.14.42.0-2.1-1.9-0.7
Delay disc. indifference point, 2s delay1.82.03.81.5-1.2-0.9-1.9
Delay disc. indifference point, 24s delay1.21.21.61.2-1.2-1.3-0.3
Delay disc. indifference point, 4s delay3.53.74.52.1-1.8-1.7-1.9
Delay disc. indifference point, 8s delay1.31.31.81.3-1.4-1.2-0.6
Indifference point function ln k0.10.10.3-0.20.30.2-0.5
Indifference point function log k0.10.10.3-0.20.30.2-0.5
Patch foraging total patch changes 0 sec1.93.13.71.9-1.8-1.8-1.6
Patch foraging total patch changes 12 sec1.31.72.11.4-1.3-1.3-1.2
Patch foraging total patch changes 18 sec1.82.33.31.7-1.8-1.8-0.3
Patch foraging total patch changes 24 sec0.91.12.0-1.00.70.81.4
Patch foraging total patch changes 6 sec1.62.32.91.6-1.6-1.7-1.2
Patch foraging time to switch 0 sec0.20.40.6-0.60.60.80.4
Patch foraging water rate 0 sec1.42.23.01.6-1.6-1.7-0.8
Patch foraging water rate 12 sec0.00.00.00.10.1-0.20.1
Patch foraging water rate 18 sec0.30.41.30.00.30.4-1.1
Patch foraging water rate 24 sec0.10.20.50.1-0.3-0.30.7
Patch foraging water rate 6 sec0.30.50.60.8-0.6-0.6-0.8
Seeking ratio, delayed vs. immediate footshock0.10.10.2-0.30.20.50.4
Distance traveled before self-admin0.90.92.30.9-0.6-0.5-1.5
Distance traveled after self-admin0.40.41.30.6-0.3-0.3-1.1
Extensor digitorum longus weight3.413.216.04.0-3.5-3.6-3.4
Bone: elastic displacement0.50.60.9-0.70.90.9-0.2
Bone: elastic work0.30.41.40.30.00.1-1.2
Bone: endosteal estimation0.10.30.40.6-0.5-0.4-0.5
Bone: endosteal perimeter0.81.62.11.5-1.4-1.2-0.9
Epididymis fat weight0.30.81.1-0.81.01.1-0.4
Delta time in closed arm before/after self-admin5.55.46.7-2.62.42.61.5
Delta time in open arm before/after self-admin9.08.711.33.4-3.1-3.0-2.2
Diff in mean of infusions in LGA sessions0.10.10.2-0.20.20.5-0.2
Delay discount exponential model param3.63.84.9-2.22.11.91.5
Extinction: sum of active levers before priming0.40.40.60.6-0.7-0.80.0
Active lever presses in extinction session 60.50.51.10.7-0.6-0.3-1.0
Bone: final force0.40.60.70.8-0.7-0.7-0.7
Bone: final moment0.60.91.11.0-0.9-0.9-1.0
Food consumed during 24 hour testing period1.41.72.91.3-1.4-1.7-0.1
Glucose14.922.433.45.0-5.6-5.8-0.2
Heart weight0.91.22.7-1.10.80.61.6
Delay discount hyperbolic model param3.73.95.0-2.22.11.91.6
Intermittent access intake day 1-15 change0.20.20.30.2-0.4-0.60.5
Intermittent access intake escalation0.10.10.20.0-0.2-0.40.5
Intermittent access intake escalation 20.10.10.10.1-0.1-0.30.4
Intermitt. access day 1 inactive lever presses1.51.62.1-1.41.20.91.4
Intermitt. access day 15 inactive lever presses0.60.71.20.7-0.9-1.10.5
Intermitt. access escalation Index0.10.10.20.1-0.1-0.30.5
Intermittent access day 1 total infusions0.10.10.30.4-0.30.3-0.5
Intermittent access day 15 total infusions0.40.40.70.8-0.8-0.5-0.4
Intermittent access terminal intake (last 3 days)0.30.30.50.7-0.7-0.4-0.4
Intermittent access total infusions0.30.40.50.7-0.7-0.3-0.6
Intermittent access day 1 locomotion0.40.40.9-0.60.50.90.3
Intermittent access day 15 total locomotion0.30.30.50.6-0.7-0.6-0.1
Intermittent access total locomotion0.50.50.80.9-0.9-0.5-0.5
Intermittent access standard deviation1.21.31.5-1.21.11.20.9
Left kidney weight0.00.10.3-0.10.20.4-0.5
Right kidney weight0.61.62.9-1.11.41.7-0.2
Cocaine induced anxiety2.42.73.61.9-1.7-1.9-1.1
Post-drug Anxiety0.70.81.21.1-1.0-1.1-0.2
Baseline Anxiety0.30.30.8-0.50.30.40.9
Tail length2.06.19.92.6-2.1-1.8-3.2
Length with tail2.59.211.63.4-3.0-2.7-3.0
Length without tail1.32.93.71.9-1.9-1.8-0.9
Times rat made contact with spout0.50.62.1-0.1-0.3-0.41.4
Lifetime Intake0.50.60.90.9-0.8-0.3-0.9
Liver weight, left1.22.03.5-1.31.61.9-0.2
Liver weight, right2.54.16.5-2.12.32.50.5
Locomotor activity0.71.21.71.3-1.3-1.2-0.1
Locomotor testing distance0.50.81.90.9-0.5-0.4-1.4
Locomotor testing rearing1.93.03.82.0-1.8-1.7-1.4
Locomotion in novel chamber3.74.66.0-2.32.32.51.2
Locomotion in novel chamber post-restriction4.04.86.9-2.31.91.82.6
Fecal boli incidents, locomotor time 10.40.40.6-0.70.60.50.7
Change in fecal boli incidents, locomotor task0.20.20.50.1-0.3-0.30.7
Time >=10cm from walls, locomotor time 10.00.00.0-0.10.10.10.2
Bouts of movement, locomotor time 10.60.60.90.7-1.0-0.90.4
Total resting periods, locomotor time 10.40.40.90.3-0.6-0.60.9
Rest time, locomotor task time 10.10.20.5-0.1-0.1-0.20.7
Distance moved, locomotor task time 10.50.61.50.7-0.4-0.3-1.2
Vertical activity count, locomotor time 10.20.30.60.5-0.3-0.2-0.8
Time >=10cm from walls, locomotor time 20.10.10.1-0.20.10.30.4
Bouts of movement, locomotor time 20.00.00.00.1-0.00.1-0.0
Total resting periods, locomotor time 20.00.00.00.1-0.10.00.0
Rest time, locomotor task time 20.10.10.2-0.40.30.20.5
Distance moved, locomotor task time 20.10.10.1-0.30.20.20.2
Vertical activity count, locomotor time 20.00.00.0-0.10.10.1-0.0
Light reinforcement 10.20.51.40.5-0.3-0.2-1.2
Light reinforcement 20.50.71.90.7-0.5-0.2-1.4
Bone: marrow area0.10.20.20.5-0.4-0.3-0.3
Bone: maximum diameter4.48.210.23.2-3.1-3.2-1.6
Bone: maximum force0.40.61.00.8-0.7-0.6-1.0
Bone: maximum moment0.61.01.61.1-0.8-0.8-1.3
Mean time between licks in bursts0.60.81.10.7-1.0-1.00.9
Mean num. licks in bursts2.93.56.4-1.91.41.42.5
Cd content in liver0.20.20.4-0.50.60.60.0
Co content in liver0.00.00.1-0.20.30.1-0.2
Cu content in liver0.30.40.60.6-0.6-0.8-0.3
Fe content in liver1.21.44.7-0.90.40.32.2
K content in liver1.31.33.80.9-0.4-0.5-2.0
Mg content in liver0.40.41.00.5-0.3-0.5-1.0
Mn content in liver0.70.82.90.30.10.0-1.7
Na content in liver3.84.05.12.2-2.2-2.3-0.9
Rb content in liver3.64.55.92.3-1.9-1.8-2.4
Se content in liver1.72.02.81.5-1.2-1.3-1.7
Sr content in liver1.61.52.21.3-1.5-1.4-0.1
Zn content in liver0.40.40.80.6-0.4-0.5-0.9
Bone: minimum diameter0.00.10.20.10.00.0-0.5
Time in familiar zone, hab. session 10.10.10.2-0.40.40.40.1
Time in novel zone, hab. session 10.60.60.90.8-0.7-0.6-0.9
Total zone transitions, hab. session 10.00.00.0-0.10.0-0.10.1
Total locomotion distance, hab. session 10.10.20.40.4-0.3-0.3-0.6
Locomotion velocity, hab. session 10.00.00.00.0-0.1-0.10.1
Time in familiar zone, hab. session 21.71.72.0-1.41.21.31.3
Time in novel zone, hab. session 22.42.42.71.7-1.4-1.6-1.5
Total zone transitions, hab. session 20.50.70.90.9-0.8-0.9-0.6
Total locomotion distance, hab. session 20.60.71.20.9-0.9-1.1-0.3
Locomotion velocity, hab. session 20.81.11.51.1-1.1-1.2-0.6
Novel to familiar place preference ratio0.90.91.41.1-1.2-1.1-0.0
Novelty place preference1.01.01.51.1-1.2-1.2-0.0
Time in novel zone, NPP test0.91.01.41.1-1.2-1.1-0.0
Total zone transitions, NPP test0.50.61.0-0.91.00.9-0.0
Total locomotion distance, NPP test0.10.10.2-0.40.50.4-0.0
Locomotion velocity, NPP test0.00.00.0-0.10.20.1-0.2
Intraocular pressure0.30.40.60.5-0.7-0.80.3
Delta distance traveled before/after self-admin0.50.50.7-0.80.70.50.9
Delta ambulatory time before/after self-admin1.01.11.4-1.21.10.81.1
Parametrial fat weight1.73.34.91.8-2.2-2.20.2
Pavlov. Cond. lever latency0.60.71.11.0-1.1-1.0-0.1
Pavlov. Cond. magazine entry latency2.53.24.2-2.12.01.81.2
Pavlov. Cond. change in total contacts0.50.60.90.8-0.7-0.9-0.5
Pavlov. Cond. index score2.73.44.5-2.12.11.91.1
Pavlov. Cond. latency score2.12.73.6-1.91.91.70.9
Pavlov. Cond. lever contacts1.21.42.0-1.41.41.20.6
Pavlov. Cond. magazine entry number3.13.95.12.3-2.1-1.8-1.6
Pavlov. Cond. intertrial magazine entries1.01.22.51.1-0.8-0.6-1.6
Pavlov. Cond. lever-magazine prob. diff.3.24.05.2-2.32.22.01.2
Pavlov. Cond. response bias1.72.12.9-1.71.71.40.8
Pavlov. Cond. lever latency4.96.69.02.8-2.2-2.2-3.0
Pavlov. Cond. magazine entry latency0.00.10.1-0.30.30.10.1
Pavlov. Cond. change in total contacts0.30.40.8-0.60.40.50.9
Pavlov. Cond. index score1.72.12.6-1.61.31.31.6
Pavlov. Cond. latency score1.72.12.7-1.61.41.31.5
Pavlov. Cond. lever contacts3.85.17.5-2.41.91.92.7
Pavlov. Cond. magazine entry number0.10.10.30.3-0.2-0.1-0.6
Pavlov. Cond. intertrial magazine entries0.00.00.0-0.0-0.0-0.00.1
Pavlov. Cond. lever-magazine prob. diff.1.51.92.4-1.51.31.31.4
Pavlov. Cond. response bias1.82.44.2-1.61.21.22.0
Time in closed arm before self-admin0.50.60.90.8-0.5-0.7-0.9
Time in closed arm after self-admin2.12.12.8-1.71.71.50.8
Time in open arm before self-admin2.72.93.6-1.91.61.71.5
Time in open arm after self-admin0.40.40.70.7-0.9-0.60.3
Bone: periosteal estimation1.22.53.21.8-1.6-1.5-1.5
Bone: periosteal perimeter2.13.64.52.1-1.9-1.8-1.6
Bone: post-yield work0.20.30.40.6-0.6-0.6-0.5
Progressive ratio test 1 active lever presses0.10.10.20.4-0.4-0.2-0.1
Progressive ratio test 1 breakpoint0.00.00.10.0-0.10.20.1
Progressive ratio test 1 inactive lever presses2.12.32.71.7-1.6-1.6-1.1
Progressive ratio test 2 active lever presses0.10.10.20.4-0.4-0.2-0.3
Progressive ratio test 2 breakpoint0.10.10.20.4-0.4-0.2-0.2
Progressive ratio test 2 inactive lever presses0.60.71.10.9-1.0-0.9-0.1
Incentive Sensitization - Responses0.00.00.10.2-0.3-0.20.1
Incentive Sensitization - Breakpoint0.00.00.00.1-0.2-0.20.2
Sum of active levers in priming session hrs 5-60.50.40.70.6-0.6-0.8-0.6
Progressive ratio0.00.00.1-0.00.10.3-0.3
Food seeking constrained by brief footshock3.13.86.72.1-1.5-1.4-2.6
Seeking ratio, punishment vs. effort1.01.21.41.2-1.0-1.1-1.2
Retroperitoneal fat weight0.00.10.4-0.30.20.10.6
Reaction time number correct0.00.10.2-0.0-0.1-0.20.5
Reaction time mean minus median2.02.64.4-1.71.31.22.1
Reaction time mean minus median AUC1.31.72.0-1.41.31.21.3
Reaction time num false alarms0.20.30.90.20.0-0.2-1.0
Reaction time num false alarms AUC0.20.31.30.10.2-0.1-1.2
Reaction time trials correct on left0.00.10.2-0.0-0.1-0.20.5
Reaction time trials on left0.10.10.3-0.1-0.0-0.20.5
Reaction time mean0.91.31.7-1.21.01.01.3
Reaction time mean AUC1.31.72.0-1.41.31.31.3
Median of all reaction times0.40.60.8-0.80.70.70.9
Reaction time omissions1.62.22.8-1.71.61.51.1
Reaction time false alarm rate0.30.51.80.10.20.0-1.4
Reaction time premature initiation rate2.83.75.1-2.12.32.20.8
Reaction time premature initiations1.62.33.1-1.71.81.60.8
Std. dev. reaction times1.72.33.8-1.61.31.21.9
Reaction time trials completed0.10.10.3-0.1-0.0-0.20.5
Reaction time trials AUC0.00.10.10.1-0.2-0.30.3
Run reversals in cocaine runway, females0.90.91.2-1.10.90.80.9
Run reversals in cocaine runway, males0.60.60.70.8-0.8-0.7-0.7
Latency to leave start box in cocaine runway0.10.10.30.10.0-0.0-0.5
Latency to leave start box in cocaine runway, F0.60.61.40.8-0.5-0.6-1.2
Latency to leave start box in cocaine runway, M0.60.60.7-0.80.70.80.6
Std. dev. time between licks in bursts0.10.10.2-0.20.1-0.00.5
Total sessions with >9 infusions0.10.10.40.10.00.2-0.6
Short access day 1 total inactive lever presses4.34.86.12.5-2.1-1.9-2.3
Short access day 10 total inactive lever presses0.50.50.7-0.90.80.60.7
Short access day 1 total infusions0.20.20.3-0.10.30.6-0.6
Short access day 10 total infusions0.10.20.40.2-0.10.4-0.6
Short access total infusions0.20.30.70.10.10.6-0.8
Short access day 1 locomotion1.01.12.1-0.91.21.4-0.3
Short access day 10 total locomotion0.30.30.7-0.40.50.8-0.2
Short access total locomotion2.12.23.9-1.51.62.00.3
One hour access (0.3 mA shock)0.10.10.20.5-0.4-0.1-0.4
Compulsive drug intake2.93.03.8-1.91.91.71.3
One hour access (shock baseline)1.41.62.31.5-1.4-1.1-1.0
Social response proportion1.21.32.51.0-1.3-1.60.5
Social responses1.51.93.41.4-1.6-1.80.0
Social time0.20.20.50.1-0.3-0.60.7
Soleus weight1.23.55.52.0-1.6-1.4-2.3
Bone: stiffness0.00.00.00.0-0.1-0.00.0
Tibialis anterior weight1.24.05.02.2-2.2-2.1-1.4
Time to tail flick, vehicle, before self-admin0.00.00.0-0.00.20.2-0.1
Time to tail flick, vehicle, after self-admin0.00.00.1-0.10.20.2-0.2
Time to tail flick, test, before self-admin0.60.60.8-0.90.80.80.5
Time to tail flick, test, after self-admin3.33.44.51.9-1.7-1.7-2.1
Delta time to tail flick, vehicle, before/after SA0.00.00.00.1-0.1-0.0-0.2
Delta time to tail flick, test, before/after SA2.01.82.11.4-1.4-1.4-1.1
Tibia length0.20.51.60.10.30.5-1.3
Ambulatory time before self-admin1.81.84.51.3-0.9-0.7-2.1
Ambulatory time after self-admin0.50.61.40.7-0.4-0.4-1.2
Bone: tissue strength0.10.10.10.3-0.4-0.30.1
Total heroin consumption0.20.20.20.5-0.4-0.3-0.5
Bone: trabecular number0.10.10.4-0.10.30.3-0.6
Bone: trabecular spacing1.01.31.71.3-1.3-1.3-0.5
Bone: trabecular thickness1.21.52.2-1.31.51.40.2
Bone: trabecular tissue density0.50.61.9-0.70.30.31.4
Water consumed over 24 hours1.01.22.01.1-1.2-1.40.1
Number of responses in last shaping day0.10.10.1-0.10.30.2-0.4