Tle5

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 Adipose alternative TSS ENSRNOT00000113477 0.04 0.03 7.2e-03 0.02 0.01 0.01 0.01 4.1e-03 1.6e-02 2.0e-02 2.0e-02
2 Adipose intron excision ratio chr7:8144252:8144373 0.03 0.02 5.6e-03 0.01 0.01 0.01 0.01 6.7e-03 2.2e-02 1.6e-02 3.4e-02
3 Brain mRNA stability ENSRNOG00000052025 0.03 0.03 7.2e-03 0.02 0.02 0.01 0.01 6.6e-03 1.0e-02 2.0e-02 1.9e-02
4 Liver intron excision ratio chr7:8144249:8144373 0.03 0.02 6.0e-03 0.02 0.01 0.02 0.02 1.8e-03 2.0e-02 5.0e-03 5.3e-03

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 strong Z-scores (below -2 or above 2): lower negative Z-scores are darker blue, higher positive Z-scores are darker red.

Trait Avg chi2 ratio Avg chi2 Max chi2 1 2 3 4
BMI with tail5.016.021.5-0.84.6-4.6-4.6
BMI without tail2.14.57.2-1.02.7-2.0-2.4
Body weight0.93.85.4-0.62.2-2.2-2.3
Epididymis fat weight0.10.30.5-0.7-0.20.70.2
Glucose0.91.41.9-0.41.4-1.3-1.4
Heart weight0.30.40.8-0.7-0.20.90.5
Left kidney weight0.20.71.71.3-0.60.10.8
Right kidney weight1.23.55.10.8-2.12.02.3
Tail length2.16.510.8-0.5-2.73.32.7
Length with tail1.55.58.20.4-2.92.62.6
Length without tail0.30.71.11.0-1.10.00.7
Liver weight, left2.23.65.4-0.82.3-1.9-2.1
Liver weight, right1.22.03.0-0.11.6-1.7-1.6
Parametrial fat weight0.30.61.0-0.4-0.91.00.8
Retroperitoneal fat weight0.51.52.4-1.01.2-1.1-1.6
Intraocular pressure0.50.72.1-1.40.60.5-0.5
Extensor digitorum longus weight3.915.822.1-2.14.6-3.9-4.7
Soleus weight6.619.126.9-0.85.0-5.2-4.9
Tibialis anterior weight0.72.33.3-0.21.7-1.8-1.7
Tibia length0.61.72.8-0.2-1.51.71.4
Number of licking bursts0.60.71.9-1.40.60.0-0.8
Food consumed during 24 hour testing period0.40.40.6-0.20.8-0.7-0.8
Times rat made contact with spout0.40.50.8-0.90.8-0.2-0.8
Mean time between licks in bursts2.73.86.3-0.5-2.12.52.1
Mean num. licks in bursts0.10.10.30.5-0.0-0.10.1
Std. dev. time between licks in bursts1.21.52.01.2-1.40.71.3
Water consumed over 24 hours0.50.61.11.10.3-1.0-0.3
Indifference point 0 sec0.10.20.30.10.3-0.6-0.5
Indifference point AUC0.70.81.40.0-0.91.21.1
Indifference point function ln k0.20.30.5-0.30.5-0.6-0.7
Indifference point function log k0.20.30.5-0.30.5-0.6-0.7
Delay discounting total patch changes 0 sec0.10.10.3-0.10.3-0.5-0.5
Delay discounting total patch changes 12 sec0.60.71.1-0.91.1-0.4-1.0
Delay discounting total patch changes 18 sec0.70.91.30.1-1.11.21.1
Delay discounting total patch changes 24 sec0.10.10.20.3-0.30.30.4
Delay discounting total patch changes 6 sec0.10.20.4-0.1-0.60.40.3
Delay discounting time to switch 0 sec1.01.63.21.01.0-1.8-1.1
Delay discounting water rate 0 sec0.40.72.1-1.50.40.5-0.5
Delay discounting water rate 12 sec1.62.43.3-1.61.8-0.8-1.7
Delay discounting water rate 18 sec1.62.16.5-2.50.90.8-0.8
Delay discounting water rate 24 sec2.94.09.8-3.11.60.1-1.9
Delay discounting water rate 6 sec0.81.23.3-1.80.01.1-0.1
Locomotor activity3.55.68.71.4-2.92.02.7
Locomotor testing distance1.93.24.80.9-2.21.62.1
Locomotor testing rearing0.00.00.1-0.30.00.20.1
Light reinforcement 14.48.011.50.5-3.43.23.1
Light reinforcement 21.41.82.90.2-1.71.51.5
Reaction time number correct1.11.52.0-1.31.4-0.6-1.4
Reaction time mean minus median0.70.91.50.5-1.01.01.2
Reaction time mean minus median AUC0.91.13.01.7-0.8-0.20.9
Reaction time num false alarms0.10.10.2-0.40.4-0.1-0.3
Reaction time num false alarms AUC0.30.40.7-0.80.7-0.1-0.6
Reaction time trials correct on left1.11.52.0-1.31.4-0.6-1.4
Reaction time trials on left1.01.41.9-1.21.4-0.6-1.3
Reaction time mean0.40.61.71.3-0.4-0.30.6
Reaction time mean AUC0.40.51.61.2-0.4-0.40.5
Median of all reaction times0.40.61.51.20.1-0.90.0
Reaction time omissions0.30.40.60.5-0.80.50.7
Reaction time false alarm rate0.00.00.00.20.0-0.10.1
Reaction time premature initiation rate0.70.91.7-1.0-0.81.30.5
Reaction time premature initiations0.81.13.2-1.8-0.11.1-0.1
Std. dev. reaction times0.50.71.01.0-0.80.41.0
Reaction time trials completed1.01.41.9-1.21.4-0.6-1.3
Reaction time trials AUC1.11.62.2-1.51.4-0.5-1.4
Social response proportion0.60.71.10.6-0.80.71.1
Social responses6.88.813.21.5-3.32.93.6
Social time0.60.81.2-0.51.0-0.9-1.1
Conditioned locomotion2.52.94.0-1.62.0-1.1-2.0
Cocaine response after cond. corrected5.56.18.7-1.82.9-2.1-2.9
Cocaine response after cond. not corrected6.06.79.4-1.93.0-2.1-3.1
Cocaine response before conditioning0.50.60.90.6-0.90.71.0
Saline control response0.10.20.2-0.50.4-0.1-0.5
Condit. Reinf. active minus inactive responses1.51.93.7-0.8-1.21.91.2
Condit. Reinf. active-inactive response ratio0.80.82.6-1.60.40.7-0.4
Condit. Reinf. active responses1.72.33.60.0-1.71.91.7
Condit. Reinf. inactive responses1.31.62.40.9-1.51.11.5
Condit. Reinf. lever presses1.52.14.0-0.6-1.42.01.4
Condit. Reinf. lever reinforcers received1.21.73.0-0.3-1.31.71.3
Pavlov. Cond. lever latency0.71.01.4-0.31.1-1.1-1.2
Pavlov. Cond. magazine entry latency0.60.71.10.6-1.00.71.0
Pavlov. Cond. change in total contacts0.70.91.4-0.0-1.01.21.1
Pavlov. Cond. index score0.81.01.50.2-1.11.11.2
Pavlov. Cond. latency score0.81.11.50.5-1.21.01.2
Pavlov. Cond. lever contacts0.60.81.3-0.1-0.91.21.0
Pavlov. Cond. magazine entry number0.91.21.7-0.61.3-1.0-1.3
Pavlov. Cond. intertrial magazine entries0.70.91.4-0.90.9-0.6-1.2
Pavlov. Cond. lever-magazine prob. diff.0.91.11.60.4-1.21.11.3
Pavlov. Cond. response bias1.31.93.0-0.1-1.41.71.5
Conditioned reinforcement - actives2.12.12.80.5-1.61.61.7
Conditioned reinforcement - inactives1.01.21.7-1.31.0-0.6-1.3
Conditioned locomotion5.97.110.30.3-3.03.23.0
Intermittent access intake day 1-15 change4.74.77.6-0.3-2.52.82.3
Intermittent access intake escalation4.34.28.1-1.3-2.02.81.7
Intermittent access intake escalation 25.15.19.9-1.1-2.23.12.1
Intermitt. access day 1 inactive lever presses0.40.40.8-0.9-0.30.90.2
Intermitt. access day 15 inactive lever presses0.30.30.5-0.6-0.50.70.4
Intermitt. access escalation Index0.70.62.01.4-0.1-0.70.3
Intermittent access day 1 total infusions0.00.00.10.10.1-0.4-0.1
Intermittent access day 15 total infusions1.21.32.3-0.7-1.21.51.1
Intermittent access terminal intake (last 3 days)0.80.91.40.1-1.01.11.2
Intermittent access total infusions0.10.10.2-0.4-0.40.40.3
Intermittent access day 1 locomotion0.20.20.3-0.4-0.50.50.4
Intermittent access day 15 total locomotion2.52.64.1-0.5-1.82.01.7
Intermittent access total locomotion0.80.91.3-0.1-1.21.11.1
Intermittent access standard deviation2.12.34.0-0.4-1.52.01.6
Cocaine induced anxiety1.31.52.01.4-1.20.71.4
Post-drug Anxiety1.41.63.71.9-1.00.31.3
Baseline Anxiety0.10.10.3-0.60.10.2-0.2
Lifetime Intake0.00.10.1-0.0-0.30.20.3
Incentive Sensitization - Responses0.20.20.5-0.4-0.20.70.3
Incentive Sensitization - Breakpoint0.20.20.6-0.1-0.30.80.5
Progressive ratio test 1 active lever presses0.20.20.3-0.50.5-0.3-0.4
Progressive ratio test 1 breakpoint0.20.20.3-0.40.6-0.4-0.4
Progressive ratio test 1 inactive lever presses0.00.00.0-0.0-0.10.00.1
Progressive ratio test 2 active lever presses0.20.20.3-0.5-0.20.60.3
Progressive ratio test 2 breakpoint0.20.30.40.1-0.50.60.6
Progressive ratio test 2 inactive lever presses0.10.10.10.2-0.30.20.3
Total sessions with >9 infusions0.40.41.21.1-0.3-0.20.7
Short access day 1 total inactive lever presses0.00.00.0-0.0-0.10.20.2
Short access day 10 total inactive lever presses0.20.20.7-0.80.00.4-0.1
Short access day 1 total infusions0.60.61.31.1-0.60.00.9
Short access day 10 total infusions0.00.00.00.10.00.00.1
Short access total infusions0.20.20.60.8-0.1-0.10.5
Short access day 1 locomotion1.01.11.90.5-1.40.71.2
Short access day 10 total locomotion2.93.15.3-0.5-2.02.31.8
Short access total locomotion0.90.91.50.3-1.10.91.2
Compulsive drug intake0.60.61.00.20.7-1.0-0.9
One hour access (0.3 mA shock)0.20.20.3-0.30.4-0.5-0.5
One hour access (shock baseline)0.10.20.6-0.7-0.00.2-0.1
Number of responses in last shaping day0.20.20.30.2-0.50.40.5
Context. condit. distance diff. score0.30.30.4-0.20.6-0.6-0.6
Locomotion velocity, session 11.31.42.01.4-1.30.61.3
Locomotion distance, session 11.01.33.41.8-0.9-0.10.9
Locomotion velocity, session 24.55.37.12.1-2.71.62.6
Locomotion distance, session 22.43.54.62.1-2.11.02.0
Locomotion velocity, session 34.45.17.50.2-2.52.72.6
Locomotion distance, session 30.00.00.1-0.20.00.20.0
Stereotopy head waving bouts, day 33.84.66.41.0-2.52.22.5
Stereotopy head waving duration, day 34.25.17.00.8-2.62.52.6
Locomotion velocity, session 70.80.81.20.6-1.10.81.0
Locomotion distance, session 70.30.30.8-0.90.5-0.0-0.5
Stereotopy head waving bouts, day 70.60.72.01.4-0.6-0.30.6
Stereotopy head waving duration, day 71.01.22.71.7-1.00.01.0
Locomotion velocity, session 81.72.04.12.0-1.40.21.4
Locomotion distance, session 80.20.30.50.7-0.50.20.5
Degree of sensitization distance0.30.30.4-0.00.5-0.7-0.6
Degree of sensitization stereotypy1.21.32.61.00.8-1.6-0.9
Condit. Reinf. active minus inactive responses3.23.54.80.4-2.22.12.1
Condit. Reinf. active-inactive response ratio1.41.52.8-0.5-1.31.71.2
Condit. Reinf. active responses3.44.15.91.3-2.41.92.3
Condit. Reinf. inactive responses1.41.62.31.5-1.40.61.3
Incentive salience index mean2.22.73.91.1-2.01.41.9
Condit. Reinf. lever presses1.11.54.12.0-1.0-0.30.9
Time in familiar zone, hab. session 11.41.32.4-0.3-1.21.51.2
Time in novel zone, hab. session 10.80.81.51.20.4-1.2-0.5
Total zone transitions, hab. session 11.42.02.61.6-1.50.81.6
Total locomotion distance, hab. session 12.63.76.42.5-2.00.72.0
Locomotion velocity, hab. session 12.43.44.32.1-2.01.02.1
Time in familiar zone, hab. session 24.14.37.9-0.8-2.12.82.1
Time in novel zone, hab. session 23.03.05.91.01.6-2.4-1.6
Total zone transitions, hab. session 20.81.03.31.8-0.3-0.80.2
Total locomotion distance, hab. session 21.21.72.11.4-1.40.81.4
Locomotion velocity, hab. session 21.52.03.21.8-1.50.71.5
Novel to familiar place preference ratio2.42.65.21.31.3-2.3-1.3
Novelty place preference1.51.63.21.30.9-1.8-0.9
Time in novel zone, NPP test1.71.73.51.21.0-1.9-1.0
Total zone transitions, NPP test0.20.30.90.9-0.2-0.30.2
Total locomotion distance, NPP test0.40.61.91.4-0.3-0.40.3
Locomotion velocity, NPP test0.40.61.81.3-0.6-0.10.6
Pavlov. Cond. lever latency0.50.61.8-1.30.50.3-0.5
Pavlov. Cond. magazine entry latency1.82.33.7-0.1-1.71.91.7
Pavlov. Cond. change in total contacts0.10.10.10.2-0.30.20.3
Pavlov. Cond. index score0.60.70.90.9-1.00.50.9
Pavlov. Cond. latency score0.81.01.71.3-1.10.41.1
Pavlov. Cond. lever contacts0.81.02.61.6-0.8-0.10.8
Pavlov. Cond. magazine entry number0.40.50.7-0.10.8-0.8-0.8
Pavlov. Cond. intertrial magazine entries0.30.30.9-1.00.20.5-0.2
Pavlov. Cond. lever-magazine prob. diff.0.81.01.40.9-1.20.81.2
Pavlov. Cond. response bias0.80.91.30.3-1.11.11.1
Bone: apparent density1.52.57.02.6-1.1-0.51.2
Bone surface3.56.215.13.9-2.2-0.22.2
Bone volume3.66.115.13.9-2.2-0.22.2
Bone: connectivity density0.81.33.92.0-0.5-0.80.5
Bone: cortical apparent density0.71.01.5-0.21.1-1.2-1.1
Bone: cortical area0.60.92.01.40.2-1.2-0.1
Bone: cortical porosity0.70.71.00.8-1.00.61.0
Bone: cortical porosity0.50.51.31.2-0.6-0.10.6
Bone: cortical thickness0.71.22.40.61.0-1.6-1.0
Bone: cortical thickness0.20.20.7-0.80.30.2-0.3
Bone: cortical tissue density0.50.71.1-0.20.9-1.0-0.9
Bone: elastic displacement0.30.30.40.2-0.60.50.6
Bone: elastic work0.10.20.30.60.1-0.6-0.1
Bone: endosteal estimation0.20.30.70.80.2-0.8-0.2
Bone: endosteal perimeter0.61.32.01.4-1.20.41.2
Bone: final force0.81.32.61.20.7-1.6-0.7
Bone: final moment0.61.02.21.10.6-1.5-0.6
Bone: marrow area0.20.30.70.70.2-0.9-0.2
Bone: maximum diameter1.32.33.11.8-1.70.71.7
Bone: maximum force0.81.32.90.90.9-1.7-0.9
Bone: maximum moment0.61.12.40.90.8-1.6-0.8
Bone: minimum diameter0.20.41.31.1-0.4-0.40.4
Bone: periosteal estimation0.51.02.51.60.0-1.2-0.0
Bone: periosteal perimeter0.71.22.91.7-1.0-0.11.0
Bone: post-yield work0.50.71.0-0.10.9-1.0-0.9
Bone: stiffness1.93.15.70.51.8-2.4-1.8
Bone: tissue strength1.21.22.20.21.1-1.5-1.1
Bone: trabecular number1.72.63.61.9-1.80.71.8
Bone: trabecular spacing2.53.26.8-2.61.7-0.1-1.7
Bone: trabecular thickness3.64.311.93.5-1.6-0.61.6
Bone: trabecular tissue density2.53.54.8-1.42.2-1.6-2.2
Breakpoint value in progressive ratio session3.53.97.0-0.4-2.02.62.1
Distance traveled before self-admin0.20.20.30.40.4-0.5-0.3
Distance traveled after self-admin0.60.71.10.4-0.90.81.1
Delta time in closed arm before/after self-admin2.52.54.9-2.21.6-0.3-1.5
Delta time in open arm before/after self-admin0.80.81.40.20.8-1.2-1.0
Diff in mean of infusions in LGA sessions1.01.11.70.0-1.01.31.3
Extinction: sum of active levers before priming1.01.02.1-1.1-0.61.50.6
Active lever presses in extinction session 62.32.23.50.5-1.51.71.9
Delta distance traveled before/after self-admin1.01.11.60.2-1.21.21.3
Delta ambulatory time before/after self-admin1.41.52.40.7-1.31.11.6
Time in closed arm before self-admin0.70.81.51.10.3-1.2-0.5
Time in closed arm after self-admin0.00.00.0-0.10.10.0-0.0
Time in open arm before self-admin0.70.81.5-0.3-0.81.20.9
Time in open arm after self-admin0.60.61.0-0.50.8-0.7-1.0
Sum of active levers in priming session hrs 5-60.00.00.1-0.20.10.0-0.1
Time to tail flick, vehicle, before self-admin1.51.62.71.7-1.20.51.4
Time to tail flick, vehicle, after self-admin0.30.30.9-0.90.30.5-0.2
Time to tail flick, test, before self-admin1.11.12.61.6-0.70.11.0
Time to tail flick, test, after self-admin0.60.61.11.1-0.60.30.9
Delta time to tail flick, vehicle, before/after SA1.71.73.1-1.81.3-0.3-1.4
Delta time to tail flick, test, before/after SA0.30.30.9-1.00.00.5-0.1
Ambulatory time before self-admin0.10.10.20.10.4-0.5-0.3
Ambulatory time after self-admin0.40.40.60.3-0.70.60.8
Total heroin consumption1.61.92.90.6-1.41.51.7
Area under the delay curve1.41.52.60.41.2-1.6-1.3
Delay disc. indifference point, 0s delay4.85.210.11.61.9-3.2-2.0
Delay disc. indifference point, 16s delay0.10.10.20.20.3-0.4-0.3
Delay disc. indifference point, 2s delay3.74.27.60.82.0-2.8-2.1
Delay disc. indifference point, 24s delay0.40.40.60.10.8-0.7-0.7
Delay disc. indifference point, 4s delay0.80.91.61.10.6-1.3-0.6
Delay disc. indifference point, 8s delay1.01.11.5-0.71.2-0.9-1.2
Delay discount exponential model param1.31.32.3-0.4-1.21.51.2
Delay discount hyperbolic model param1.81.93.3-0.4-1.41.81.5
Fecal boli incidents, locomotor time 12.12.23.0-1.21.7-1.2-1.7
Change in fecal boli incidents, locomotor task1.21.12.21.5-1.00.31.1
Time >=10cm from walls, locomotor time 10.40.41.11.00.1-0.8-0.2
Bouts of movement, locomotor time 10.10.10.2-0.2-0.30.40.4
Total resting periods, locomotor time 10.20.20.30.1-0.50.50.6
Rest time, locomotor task time 10.20.20.30.5-0.50.30.6
Distance moved, locomotor task time 10.40.41.11.00.1-0.8-0.2
Vertical activity count, locomotor time 10.10.10.2-0.40.3-0.1-0.4
Time >=10cm from walls, locomotor time 21.11.21.7-0.71.2-1.0-1.3
Bouts of movement, locomotor time 20.80.91.41.2-1.00.41.0
Total resting periods, locomotor time 20.91.01.81.3-1.00.31.0
Rest time, locomotor task time 20.80.91.21.0-1.00.61.1
Distance moved, locomotor task time 21.01.11.6-1.01.1-0.8-1.3
Vertical activity count, locomotor time 21.41.62.2-1.51.3-0.6-1.4
Weight adjusted by age0.60.61.4-0.71.2-0.4-0.8
Seeking ratio, delayed vs. immediate footshock4.55.17.70.9-2.82.32.5
Locomotion in novel chamber0.20.30.8-0.1-0.20.90.5
Locomotion in novel chamber post-restriction0.50.60.80.3-0.70.90.9
Progressive ratio0.30.40.6-0.60.8-0.3-0.7
Food seeking constrained by brief footshock0.10.10.10.0-0.30.40.3
Seeking ratio, punishment vs. effort0.10.20.40.3-0.60.20.5
Run reversals in cocaine runway, females0.80.81.20.9-0.90.71.1
Run reversals in cocaine runway, males1.31.32.10.9-1.11.01.5
Latency to leave start box in cocaine runway1.82.02.61.6-1.60.71.6
Latency to leave start box in cocaine runway, F1.51.74.92.2-0.8-0.41.0
Latency to leave start box in cocaine runway, M1.91.93.6-1.0-1.21.91.2
Cd content in liver0.10.10.20.10.3-0.5-0.3
Co content in liver0.40.40.9-0.5-0.61.00.5
Cu content in liver0.40.41.31.2-0.3-0.50.2
Fe content in liver0.10.20.5-0.70.20.2-0.2
K content in liver0.50.51.31.1-0.5-0.20.5
Mg content in liver0.70.72.11.4-0.6-0.20.7
Mn content in liver3.33.76.92.6-1.90.52.0
Na content in liver0.60.71.40.60.7-1.2-0.7
Rb content in liver0.60.72.01.40.0-0.90.1
Se content in liver0.20.20.30.3-0.50.40.5
Sr content in liver0.60.61.8-1.30.50.3-0.5
Zn content in liver1.71.83.6-1.7-0.61.90.7