Zkscan2

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 Zkscan2 0.04 0.03 6.2e-05 0.04 0.04 0.03 0.04 3.2e-05 6.5e-05 1.9e-04 5.8e-05
2 Brain gene expression Zkscan2 0.1 0.05 8.5e-09 0.09 0.08 0.08 0.08 2.0e-08 2.8e-08 6.3e-08 3.3e-08
3 Brain isoform ratio XM_039101194.2 0.06 0.04 1.1e-04 0.01 0.05 0.04 0.04 1.4e-02 2.4e-05 6.2e-05 1.5e-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
Bone: apparent density0.30.60.91.00.80.3
Area under the delay curve0.10.20.6-0.1-0.10.7
BMI with tail3.211.315.43.83.92.0
BMI without tail2.513.519.44.24.41.7
Body weight2.912.518.94.34.30.1
Weight adjusted by age0.30.81.20.90.71.1
Bone surface0.71.52.31.51.30.7
Bone volume1.02.13.01.71.60.8
Breakpoint value in progressive ratio session0.20.30.70.0-0.4-0.8
Number of licking bursts2.47.48.7-2.9-2.22.9
Context. condit. distance diff. score0.60.81.8-0.5-0.51.3
Locomotion velocity, session 10.10.20.30.5-0.20.4
Locomotion distance, session 10.10.20.30.60.30.3
Locomotion velocity, session 20.50.71.10.90.21.0
Locomotion distance, session 20.40.60.80.90.90.4
Locomotion velocity, session 31.21.32.41.21.60.1
Locomotion distance, session 30.10.20.3-0.20.40.5
Stereotopy head waving bouts, day 30.50.61.80.30.0-1.4
Stereotopy head waving duration, day 30.30.30.60.60.3-0.8
Locomotion velocity, session 70.60.61.60.40.1-1.3
Locomotion distance, session 70.81.01.4-0.9-1.21.0
Stereotopy head waving bouts, day 70.30.40.50.30.7-0.7
Stereotopy head waving duration, day 71.01.31.91.21.4-0.7
Locomotion velocity, session 80.10.20.30.40.50.2
Locomotion distance, session 80.50.72.2-0.0-0.11.5
Degree of sensitization distance0.60.71.3-0.7-1.10.4
Degree of sensitization stereotypy0.40.40.90.40.9-0.5
Conditioned locomotion0.10.10.2-0.2-0.4-0.4
Cocaine response after cond. corrected0.71.01.4-1.2-1.10.6
Cocaine response after cond. not corrected0.00.00.00.10.2-0.2
Cocaine response before conditioning4.25.06.92.62.6-1.2
Saline control response0.30.40.7-0.4-0.40.9
Bone: connectivity density0.10.10.20.30.30.5
Bone: cortical apparent density0.71.11.3-1.1-1.10.9
Bone: cortical area3.15.16.42.52.51.7
Bone: cortical porosity1.92.03.41.41.8-0.8
Bone: cortical porosity1.71.82.31.11.5-1.4
Bone: cortical thickness1.72.84.31.21.62.1
Bone: cortical thickness0.91.12.90.20.61.7
Bone: cortical tissue density0.71.11.4-1.2-1.10.8
Condit. Reinf. active minus inactive responses0.10.10.2-0.1-0.40.2
Condit. Reinf. active-inactive response ratio0.40.41.20.3-0.0-1.1
Condit. Reinf. active responses0.70.92.4-0.3-0.61.5
Condit. Reinf. inactive responses1.92.57.1-0.4-0.62.7
Incentive salience index mean0.20.30.40.60.30.6
Condit. Reinf. lever presses0.91.32.01.10.91.4
Condit. Reinf. active minus inactive responses0.50.82.20.2-0.01.5
Condit. Reinf. active-inactive response ratio1.82.88.0-0.6-0.42.8
Condit. Reinf. active responses0.30.51.3-0.1-0.11.2
Condit. Reinf. inactive responses0.40.61.30.50.5-1.2
Condit. Reinf. lever presses0.81.11.21.01.11.1
Condit. Reinf. lever reinforcers received0.40.61.5-0.3-0.31.2
Conditioned reinforcement - actives0.10.50.8-0.6-0.6-0.9
Conditioned reinforcement - inactives0.20.51.3-0.5-1.10.2
Conditioned locomotion0.71.01.5-1.2-1.0-0.9
Patch foraging indifference point 0 sec0.30.81.31.11.00.2
Patch foraging indifference point AUC0.61.42.71.21.60.2
Delay disc. indifference point, 0s delay0.30.40.60.80.70.1
Delay disc. indifference point, 16s delay1.11.51.7-1.3-1.31.0
Delay disc. indifference point, 2s delay2.93.25.02.22.10.4
Delay disc. indifference point, 24s delay0.30.50.7-0.8-0.80.3
Delay disc. indifference point, 4s delay0.40.41.3-0.0-0.01.1
Delay disc. indifference point, 8s delay0.00.00.10.00.00.3
Indifference point function ln k0.41.21.41.21.01.0
Indifference point function log k0.41.21.41.21.01.0
Patch foraging total patch changes 0 sec0.30.91.41.21.00.5
Patch foraging total patch changes 12 sec0.00.00.0-0.1-0.1-0.1
Patch foraging total patch changes 18 sec0.00.00.1-0.3-0.10.1
Patch foraging total patch changes 24 sec0.00.10.40.00.10.6
Patch foraging total patch changes 6 sec0.10.20.40.30.60.5
Patch foraging time to switch 0 sec1.03.75.9-2.1-2.4-0.8
Patch foraging water rate 0 sec0.31.11.51.21.10.8
Patch foraging water rate 12 sec0.41.63.70.40.91.9
Patch foraging water rate 18 sec0.51.11.71.01.30.7
Patch foraging water rate 24 sec1.11.62.71.01.01.6
Patch foraging water rate 6 sec0.60.82.50.00.31.6
Seeking ratio, delayed vs. immediate footshock0.10.20.3-0.5-0.5-0.3
Distance traveled before self-admin0.20.50.8-0.8-0.9-0.2
Distance traveled after self-admin0.51.01.7-1.0-1.3-0.5
Extensor digitorum longus weight4.518.029.05.05.4-0.8
Bone: elastic displacement1.01.33.3-0.6-0.4-1.8
Bone: elastic work0.10.20.30.50.4-0.3
Bone: endosteal estimation1.12.84.32.12.00.2
Bone: endosteal perimeter0.71.73.11.81.4-0.3
Epididymis fat weight0.52.73.92.01.80.9
Delta time in closed arm before/after self-admin0.30.20.3-0.6-0.6-0.3
Delta time in open arm before/after self-admin2.02.43.41.81.80.8
Diff in mean of infusions in LGA sessions0.20.41.3-0.1-0.11.1
Delay discount exponential model param0.10.10.40.20.2-0.6
Extinction: sum of active levers before priming0.20.40.4-0.5-0.7-0.6
Active lever presses in extinction session 61.31.32.21.51.3-0.2
Bone: final force4.06.59.33.02.91.4
Bone: final moment3.55.78.42.92.61.3
Food consumed during 24 hour testing period0.71.42.21.11.50.9
Glucose0.40.70.90.80.9-0.6
Heart weight0.81.22.61.61.00.1
Delay discount hyperbolic model param0.10.20.4-0.2-0.1-0.7
Intermittent access intake day 1-15 change0.21.01.4-1.0-1.20.8
Intermittent access intake escalation0.20.81.1-0.8-1.10.9
Intermittent access intake escalation 20.50.81.2-0.8-1.10.7
Intermitt. access day 1 inactive lever presses0.11.52.1-1.2-1.01.5
Intermitt. access day 15 inactive lever presses0.10.50.60.80.70.6
Intermitt. access escalation Index0.00.00.1-0.20.20.3
Intermittent access day 1 total infusions0.00.10.2-0.50.30.3
Intermittent access day 15 total infusions0.61.41.9-1.4-0.81.2
Intermittent access terminal intake (last 3 days)0.30.50.7-0.8-0.40.8
Intermittent access total infusions0.20.20.3-0.40.40.6
Intermittent access day 1 locomotion0.30.61.4-0.70.01.2
Intermittent access day 15 total locomotion0.41.63.6-1.0-0.61.9
Intermittent access total locomotion0.60.81.8-0.30.71.3
Intermittent access standard deviation0.10.51.20.50.31.1
Left kidney weight2.217.927.35.05.2-0.9
Right kidney weight2.215.223.84.64.9-0.7
Cocaine induced anxiety0.20.60.9-1.0-0.90.5
Post-drug Anxiety0.00.00.10.1-0.0-0.3
Baseline Anxiety1.01.72.41.51.4-0.8
Tail length0.73.96.32.52.2-0.7
Length with tail0.93.85.21.81.8-2.3
Length without tail0.42.15.60.50.7-2.4
Times rat made contact with spout0.40.51.2-0.40.41.1
Lifetime Intake0.10.10.3-0.30.50.1
Liver weight, left0.40.71.50.40.7-1.2
Liver weight, right0.61.02.60.50.3-1.6
Locomotor activity1.11.74.20.80.5-2.0
Locomotor testing distance0.10.20.20.40.5-0.5
Locomotor testing rearing0.31.22.0-1.4-1.2-0.2
Locomotion in novel chamber0.71.02.50.30.61.6
Locomotion in novel chamber post-restriction1.03.03.7-1.9-1.61.7
Fecal boli incidents, locomotor time 10.51.42.2-0.9-1.1-1.5
Change in fecal boli incidents, locomotor task1.01.11.80.91.30.8
Time >=10cm from walls, locomotor time 10.30.40.6-0.6-0.7-0.6
Bouts of movement, locomotor time 10.10.10.2-0.3-0.3-0.4
Total resting periods, locomotor time 10.00.10.1-0.2-0.2-0.3
Rest time, locomotor task time 10.30.40.6-0.8-0.6-0.6
Distance moved, locomotor task time 10.51.53.40.90.51.9
Vertical activity count, locomotor time 10.10.40.70.70.20.8
Time >=10cm from walls, locomotor time 20.10.30.70.20.2-0.8
Bouts of movement, locomotor time 20.31.01.7-0.7-0.91.3
Total resting periods, locomotor time 20.20.61.2-0.4-0.61.1
Rest time, locomotor task time 20.40.60.90.30.9-1.0
Distance moved, locomotor task time 21.31.83.9-0.5-1.02.0
Vertical activity count, locomotor time 20.40.92.0-0.8-1.40.5
Light reinforcement 10.41.93.3-1.2-1.0-1.8
Light reinforcement 20.10.10.30.50.2-0.4
Bone: marrow area1.12.84.22.02.00.3
Bone: maximum diameter2.24.56.62.52.60.8
Bone: maximum force3.05.98.83.02.81.1
Bone: maximum moment2.75.17.82.82.51.1
Mean time between licks in bursts0.42.32.9-1.7-1.1-1.7
Mean num. licks in bursts1.74.65.12.22.0-2.3
Cd content in liver0.60.71.1-1.0-0.90.3
Co content in liver1.21.42.51.61.20.5
Cu content in liver1.31.62.41.11.11.5
Fe content in liver0.40.40.7-0.7-0.40.8
K content in liver2.42.43.71.91.80.4
Mg content in liver1.41.52.21.51.1-1.0
Mn content in liver0.70.81.31.11.1-0.1
Na content in liver3.13.55.32.32.2-0.7
Rb content in liver0.30.41.0-0.40.31.0
Se content in liver0.30.30.90.1-0.20.9
Sr content in liver0.40.40.90.30.90.4
Zn content in liver0.81.01.21.11.10.8
Bone: minimum diameter1.22.33.41.81.70.8
Time in familiar zone, hab. session 11.31.44.1-0.2-0.02.0
Time in novel zone, hab. session 12.62.98.60.1-0.1-2.9
Total zone transitions, hab. session 11.21.73.01.51.70.1
Total locomotion distance, hab. session 11.92.74.02.01.9-0.5
Locomotion velocity, hab. session 11.42.03.01.61.7-0.5
Time in familiar zone, hab. session 21.51.73.90.80.82.0
Time in novel zone, hab. session 21.01.12.2-0.6-0.8-1.5
Total zone transitions, hab. session 20.81.01.41.01.2-0.9
Total locomotion distance, hab. session 20.70.91.11.01.1-0.7
Locomotion velocity, hab. session 21.01.42.01.31.4-0.7
Novel to familiar place preference ratio1.31.73.4-1.1-1.8-0.7
Novelty place preference1.21.53.3-1.0-1.8-0.4
Time in novel zone, NPP test1.41.73.7-1.1-1.9-0.3
Total zone transitions, NPP test1.42.03.61.61.90.1
Total locomotion distance, NPP test0.91.21.91.31.4-0.2
Locomotion velocity, NPP test1.21.72.71.51.6-0.4
Intraocular pressure0.81.42.5-1.6-1.2-0.5
Delta distance traveled before/after self-admin0.30.40.6-0.8-0.70.1
Delta ambulatory time before/after self-admin0.30.50.8-0.9-0.80.3
Parametrial fat weight5.914.020.94.64.21.9
Pavlov. Cond. lever latency0.91.11.7-1.3-1.30.2
Pavlov. Cond. magazine entry latency1.62.02.21.51.3-1.5
Pavlov. Cond. change in total contacts0.50.50.80.60.7-0.9
Pavlov. Cond. index score0.70.91.21.11.1-0.5
Pavlov. Cond. latency score1.21.52.21.51.4-0.5
Pavlov. Cond. lever contacts1.21.42.11.51.4-0.6
Pavlov. Cond. magazine entry number0.70.91.6-0.8-0.61.3
Pavlov. Cond. intertrial magazine entries0.80.91.5-0.7-0.91.2
Pavlov. Cond. lever-magazine prob. diff.0.70.91.31.11.1-0.5
Pavlov. Cond. response bias0.60.71.10.80.6-1.1
Pavlov. Cond. lever latency0.91.31.8-1.0-1.4-1.1
Pavlov. Cond. magazine entry latency0.81.11.6-1.3-1.1-0.6
Pavlov. Cond. change in total contacts0.10.20.50.20.20.7
Pavlov. Cond. index score0.10.10.3-0.10.00.6
Pavlov. Cond. latency score0.00.10.20.10.20.4
Pavlov. Cond. lever contacts0.91.32.01.01.40.9
Pavlov. Cond. magazine entry number1.51.93.31.81.5-0.5
Pavlov. Cond. intertrial magazine entries1.94.26.62.62.4-0.5
Pavlov. Cond. lever-magazine prob. diff.0.10.10.1-0.3-0.20.3
Pavlov. Cond. response bias0.40.51.4-0.50.11.2
Time in closed arm before self-admin0.30.50.6-0.7-0.80.5
Time in closed arm after self-admin2.52.84.6-2.2-1.90.5
Time in open arm before self-admin0.30.41.1-0.10.0-1.0
Time in open arm after self-admin2.93.25.12.32.10.2
Bone: periosteal estimation3.07.19.83.13.01.6
Bone: periosteal perimeter1.32.74.32.11.9-0.1
Bone: post-yield work3.66.210.02.93.20.4
Progressive ratio test 1 active lever presses0.10.10.3-0.50.0-0.2
Progressive ratio test 1 breakpoint0.10.10.4-0.6-0.0-0.2
Progressive ratio test 1 inactive lever presses0.00.00.00.10.2-0.2
Progressive ratio test 2 active lever presses1.11.53.2-1.8-1.1-0.1
Progressive ratio test 2 breakpoint1.01.32.8-1.7-1.0-0.1
Progressive ratio test 2 inactive lever presses0.10.20.3-0.3-0.2-0.6
Incentive Sensitization - Responses0.30.70.8-0.9-0.9-0.8
Incentive Sensitization - Breakpoint0.20.50.6-0.7-0.6-0.8
Sum of active levers in priming session hrs 5-61.71.62.01.11.2-1.4
Progressive ratio0.30.92.10.60.5-1.4
Food seeking constrained by brief footshock0.10.20.50.1-0.1-0.7
Seeking ratio, punishment vs. effort0.00.00.10.10.00.3
Retroperitoneal fat weight1.36.08.82.93.0-0.8
Reaction time number correct0.51.63.0-1.7-1.3-0.4
Reaction time mean minus median0.00.10.20.30.4-0.1
Reaction time mean minus median AUC0.00.10.1-0.4-0.2-0.1
Reaction time num false alarms0.20.30.5-0.3-0.70.4
Reaction time num false alarms AUC0.00.00.00.1-0.20.1
Reaction time trials correct on left0.51.63.0-1.7-1.3-0.4
Reaction time trials on left0.51.62.9-1.7-1.3-0.4
Reaction time mean0.51.13.10.20.3-1.7
Reaction time mean AUC0.51.23.5-0.00.1-1.9
Median of all reaction times0.91.44.10.10.2-2.0
Reaction time omissions0.50.82.20.10.5-1.5
Reaction time false alarm rate0.20.20.50.70.10.4
Reaction time premature initiation rate0.71.03.0-0.3-0.1-1.7
Reaction time premature initiations0.72.04.0-1.2-0.8-2.0
Std. dev. reaction times0.10.30.70.40.4-0.8
Reaction time trials completed0.51.62.9-1.7-1.3-0.4
Reaction time trials AUC0.41.32.3-1.5-1.2-0.4
Run reversals in cocaine runway, females1.61.82.81.71.4-0.9
Run reversals in cocaine runway, males0.10.10.2-0.4-0.50.2
Latency to leave start box in cocaine runway1.01.42.60.81.01.6
Latency to leave start box in cocaine runway, F0.70.82.2-0.6-0.01.5
Latency to leave start box in cocaine runway, M1.81.72.81.71.30.7
Std. dev. time between licks in bursts0.10.10.2-0.40.0-0.5
Total sessions with >9 infusions0.30.40.90.40.9-0.4
Short access day 1 total inactive lever presses0.30.71.9-0.00.2-1.4
Short access day 10 total inactive lever presses0.11.01.9-1.4-1.00.5
Short access day 1 total infusions0.91.82.9-1.3-0.9-1.7
Short access day 10 total infusions0.10.20.4-0.70.10.1
Short access total infusions0.20.20.4-0.20.5-0.6
Short access day 1 locomotion0.91.63.3-1.8-1.20.1
Short access day 10 total locomotion0.20.72.10.30.31.4
Short access total locomotion0.40.60.80.40.90.9
One hour access (0.3 mA shock)0.10.10.20.40.2-0.0
Compulsive drug intake1.01.52.91.71.0-0.7
One hour access (shock baseline)0.10.40.7-0.8-0.30.6
Social response proportion0.50.61.0-0.6-1.0-0.7
Social responses0.20.30.4-0.5-0.6-0.6
Social time0.60.91.1-0.9-0.9-1.0
Soleus weight3.610.515.94.03.81.1
Bone: stiffness4.48.010.93.33.02.0
Tibialis anterior weight6.622.637.15.56.1-0.3
Time to tail flick, vehicle, before self-admin1.01.21.91.41.2-0.4
Time to tail flick, vehicle, after self-admin0.00.10.20.40.1-0.3
Time to tail flick, test, before self-admin1.31.52.71.11.7-0.6
Time to tail flick, test, after self-admin0.80.92.3-0.4-0.31.5
Delta time to tail flick, vehicle, before/after SA0.10.10.30.10.10.5
Delta time to tail flick, test, before/after SA0.50.71.5-0.3-0.61.2
Tibia length0.10.20.40.60.50.0
Ambulatory time before self-admin0.41.11.7-1.2-1.30.5
Ambulatory time after self-admin1.22.03.1-1.7-1.8-0.2
Bone: tissue strength0.20.20.40.20.3-0.6
Total heroin consumption0.80.92.10.70.4-1.4
Bone: trabecular number0.20.40.50.50.70.5
Bone: trabecular spacing0.71.02.6-0.4-0.5-1.6
Bone: trabecular thickness1.01.42.01.41.20.9
Bone: trabecular tissue density0.61.11.91.41.0-0.7
Water consumed over 24 hours0.71.32.2-1.5-1.10.6
Number of responses in last shaping day0.91.22.9-0.8-0.2-1.7