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 5.7e-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 6.2e-08 3.5e-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 density0.30.60.9-1.0-0.80.3
Area under the delay curve0.20.20.60.10.10.7
BMI with tail3.311.315.4-3.8-3.92.0
BMI without tail4.413.519.4-4.2-4.41.7
Body weight2.912.518.9-4.3-4.30.1
Weight adjusted by age0.50.81.2-0.9-0.71.1
Bone surface0.81.52.3-1.5-1.30.7
Bone volume1.12.13.0-1.7-1.60.8
Breakpoint value in progressive ratio session0.20.30.7-0.00.4-0.8
Number of licking bursts4.67.48.72.92.22.9
Context. condit. distance diff. score0.60.81.80.50.51.3
Locomotion velocity, session 10.20.20.3-0.50.20.4
Locomotion distance, session 10.10.20.3-0.6-0.30.3
Locomotion velocity, session 20.60.71.1-0.9-0.21.0
Locomotion distance, session 20.40.60.8-0.9-0.90.4
Locomotion velocity, session 31.21.32.4-1.2-1.60.1
Locomotion distance, session 30.20.20.30.2-0.40.5
Stereotopy head waving bouts, day 30.60.61.8-0.3-0.0-1.4
Stereotopy head waving duration, day 30.30.30.6-0.6-0.3-0.8
Locomotion velocity, session 70.60.61.6-0.4-0.1-1.3
Locomotion distance, session 70.81.01.40.91.21.0
Stereotopy head waving bouts, day 70.30.40.5-0.3-0.7-0.7
Stereotopy head waving duration, day 71.11.31.9-1.2-1.4-0.7
Locomotion velocity, session 80.10.20.3-0.4-0.50.2
Locomotion distance, session 80.50.72.20.00.11.5
Degree of sensitization distance0.60.71.30.71.10.4
Degree of sensitization stereotypy0.40.40.9-0.4-0.9-0.5
Conditioned locomotion0.10.10.20.20.4-0.4
Cocaine response after cond. corrected0.81.01.41.21.10.6
Cocaine response after cond. not corrected0.00.00.0-0.1-0.2-0.2
Cocaine response before conditioning4.25.06.9-2.6-2.6-1.2
Saline control response0.30.40.70.40.40.9
Bone: connectivity density0.10.10.2-0.3-0.30.5
Bone: cortical apparent density0.81.11.31.11.10.9
Bone: cortical area3.35.16.4-2.5-2.51.7
Bone: cortical porosity1.92.03.4-1.4-1.8-0.8
Bone: cortical porosity1.71.82.3-1.1-1.5-1.4
Bone: cortical thickness1.72.84.3-1.2-1.62.1
Bone: cortical thickness0.91.12.9-0.2-0.61.7
Bone: cortical tissue density0.81.11.41.21.10.8
Condit. Reinf. active minus inactive responses0.10.10.20.10.40.2
Condit. Reinf. active-inactive response ratio0.40.41.2-0.30.0-1.1
Condit. Reinf. active responses0.80.92.40.30.61.5
Condit. Reinf. inactive responses2.12.57.10.40.62.7
Incentive salience index mean0.20.30.4-0.6-0.30.6
Condit. Reinf. lever presses1.01.32.0-1.1-0.91.4
Condit. Reinf. active minus inactive responses0.60.82.2-0.20.01.5
Condit. Reinf. active-inactive response ratio2.42.88.00.60.42.8
Condit. Reinf. active responses0.30.51.30.10.11.2
Condit. Reinf. inactive responses0.50.61.3-0.5-0.5-1.2
Condit. Reinf. lever presses0.81.11.2-1.0-1.11.1
Condit. Reinf. lever reinforcers received0.40.61.50.30.31.2
Conditioned reinforcement - actives0.30.50.80.60.6-0.9
Conditioned reinforcement - inactives0.40.51.30.51.10.2
Conditioned locomotion0.81.01.51.21.0-0.9
Patch foraging indifference point 0 sec0.40.81.3-1.1-1.00.2
Patch foraging indifference point AUC1.01.42.7-1.2-1.60.2
Delay disc. indifference point, 0s delay0.30.40.6-0.8-0.70.1
Delay disc. indifference point, 16s delay1.41.51.71.31.31.0
Delay disc. indifference point, 2s delay2.93.25.0-2.2-2.10.4
Delay disc. indifference point, 24s delay0.40.50.70.80.80.3
Delay disc. indifference point, 4s delay0.40.41.30.00.01.1
Delay disc. indifference point, 8s delay0.00.00.1-0.0-0.00.3
Indifference point function ln k0.71.21.4-1.2-1.01.0
Indifference point function log k0.71.21.4-1.2-1.01.0
Patch foraging total patch changes 0 sec0.50.91.4-1.2-1.00.5
Patch foraging total patch changes 12 sec0.00.00.00.10.1-0.1
Patch foraging total patch changes 18 sec0.00.00.10.30.10.1
Patch foraging total patch changes 24 sec0.10.10.4-0.0-0.10.6
Patch foraging total patch changes 6 sec0.20.20.4-0.3-0.60.5
Patch foraging time to switch 0 sec1.83.75.92.12.4-0.8
Patch foraging water rate 0 sec0.51.11.5-1.2-1.10.8
Patch foraging water rate 12 sec0.81.63.7-0.4-0.91.9
Patch foraging water rate 18 sec0.71.11.7-1.0-1.30.7
Patch foraging water rate 24 sec1.11.62.7-1.0-1.01.6
Patch foraging water rate 6 sec0.60.82.5-0.0-0.31.6
Seeking ratio, delayed vs. immediate footshock0.10.20.30.50.5-0.3
Distance traveled before self-admin0.40.50.80.80.9-0.2
Distance traveled after self-admin0.81.01.71.01.3-0.5
Extensor digitorum longus weight4.618.029.0-5.0-5.4-0.8
Bone: elastic displacement1.11.33.30.60.4-1.8
Bone: elastic work0.10.20.3-0.5-0.4-0.3
Bone: endosteal estimation1.32.84.3-2.1-2.00.2
Bone: endosteal perimeter0.81.73.1-1.8-1.4-0.3
Epididymis fat weight0.82.73.9-2.0-1.80.9
Delta time in closed arm before/after self-admin0.30.20.30.60.6-0.3
Delta time in open arm before/after self-admin2.42.43.4-1.8-1.80.8
Diff in mean of infusions in LGA sessions0.40.41.30.10.11.1
Delay discount exponential model param0.10.10.4-0.2-0.2-0.6
Extinction: sum of active levers before priming0.30.40.40.50.7-0.6
Active lever presses in extinction session 61.31.32.2-1.5-1.3-0.2
Bone: final force4.16.59.3-3.0-2.91.4
Bone: final moment3.55.78.4-2.9-2.61.3
Food consumed during 24 hour testing period1.01.42.2-1.1-1.50.9
Glucose0.40.70.9-0.8-0.9-0.6
Heart weight0.81.22.6-1.6-1.00.1
Delay discount hyperbolic model param0.10.20.40.20.1-0.7
Intermittent access intake day 1-15 change0.51.01.41.01.20.8
Intermittent access intake escalation0.40.81.10.81.10.9
Intermittent access intake escalation 20.70.81.20.81.10.7
Intermitt. access day 1 inactive lever presses0.41.52.11.21.01.5
Intermitt. access day 15 inactive lever presses0.20.50.6-0.8-0.70.6
Intermitt. access escalation Index0.00.00.10.2-0.20.3
Intermittent access day 1 total infusions0.10.10.20.5-0.30.3
Intermittent access day 15 total infusions0.91.41.91.40.81.2
Intermittent access terminal intake (last 3 days)0.40.50.70.80.40.8
Intermittent access total infusions0.20.20.30.4-0.40.6
Intermittent access day 1 locomotion0.50.61.40.7-0.01.2
Intermittent access day 15 total locomotion0.91.63.61.00.61.9
Intermittent access total locomotion0.60.81.80.3-0.71.3
Intermittent access standard deviation0.20.51.2-0.5-0.31.1
Left kidney weight3.817.927.3-5.0-5.2-0.9
Right kidney weight3.315.223.8-4.6-4.9-0.7
Cocaine induced anxiety0.50.60.91.00.90.5
Post-drug Anxiety0.00.00.1-0.10.0-0.3
Baseline Anxiety1.41.72.4-1.5-1.4-0.8
Tail length1.03.96.3-2.5-2.2-0.7
Length with tail0.93.85.2-1.8-1.8-2.3
Length without tail0.72.15.6-0.5-0.7-2.4
Times rat made contact with spout0.40.51.20.4-0.41.1
Lifetime Intake0.10.10.30.3-0.50.1
Liver weight, left0.40.71.5-0.4-0.7-1.2
Liver weight, right0.61.02.6-0.5-0.3-1.6
Locomotor activity1.11.74.2-0.8-0.5-2.0
Locomotor testing distance0.10.20.2-0.4-0.5-0.5
Locomotor testing rearing0.61.22.01.41.2-0.2
Locomotion in novel chamber0.81.02.5-0.3-0.61.6
Locomotion in novel chamber post-restriction1.83.03.71.91.61.7
Fecal boli incidents, locomotor time 11.01.42.20.91.1-1.5
Change in fecal boli incidents, locomotor task1.11.11.8-0.9-1.30.8
Time >=10cm from walls, locomotor time 10.40.40.60.60.7-0.6
Bouts of movement, locomotor time 10.10.10.20.30.3-0.4
Total resting periods, locomotor time 10.10.10.10.20.2-0.3
Rest time, locomotor task time 10.30.40.60.80.6-0.6
Distance moved, locomotor task time 11.01.53.4-0.9-0.51.9
Vertical activity count, locomotor time 10.20.40.7-0.7-0.20.8
Time >=10cm from walls, locomotor time 20.20.30.7-0.2-0.2-0.8
Bouts of movement, locomotor time 20.61.01.70.70.91.3
Total resting periods, locomotor time 20.30.61.20.40.61.1
Rest time, locomotor task time 20.50.60.9-0.3-0.9-1.0
Distance moved, locomotor task time 21.51.83.90.51.02.0
Vertical activity count, locomotor time 20.70.92.00.81.40.5
Light reinforcement 10.81.93.31.21.0-1.8
Light reinforcement 20.10.10.3-0.5-0.2-0.4
Bone: marrow area1.32.84.2-2.0-2.00.3
Bone: maximum diameter2.44.56.6-2.5-2.60.8
Bone: maximum force3.35.98.8-3.0-2.81.1
Bone: maximum moment2.95.17.8-2.8-2.51.1
Mean time between licks in bursts0.92.32.91.71.1-1.7
Mean num. licks in bursts3.04.65.1-2.2-2.0-2.3
Cd content in liver0.60.71.11.00.90.3
Co content in liver1.21.42.5-1.6-1.20.5
Cu content in liver1.31.62.4-1.1-1.11.5
Fe content in liver0.40.40.70.70.40.8
K content in liver2.52.43.7-1.9-1.80.4
Mg content in liver1.51.52.2-1.5-1.1-1.0
Mn content in liver0.80.81.3-1.1-1.1-0.1
Na content in liver3.33.55.3-2.3-2.2-0.7
Rb content in liver0.30.41.00.4-0.31.0
Se content in liver0.30.30.9-0.10.20.9
Sr content in liver0.40.40.9-0.3-0.90.4
Zn content in liver0.91.01.2-1.1-1.10.8
Bone: minimum diameter1.32.33.4-1.8-1.70.8
Time in familiar zone, hab. session 11.41.44.10.20.02.0
Time in novel zone, hab. session 12.92.98.6-0.10.1-2.9
Total zone transitions, hab. session 11.21.73.0-1.5-1.70.1
Total locomotion distance, hab. session 11.92.74.0-2.0-1.9-0.5
Locomotion velocity, hab. session 11.42.03.0-1.6-1.7-0.5
Time in familiar zone, hab. session 21.61.73.9-0.8-0.82.0
Time in novel zone, hab. session 21.11.12.20.60.8-1.5
Total zone transitions, hab. session 20.81.01.4-1.0-1.2-0.9
Total locomotion distance, hab. session 20.70.91.1-1.0-1.1-0.7
Locomotion velocity, hab. session 21.01.42.0-1.3-1.4-0.7
Novel to familiar place preference ratio1.51.73.41.11.8-0.7
Novelty place preference1.41.53.31.01.8-0.4
Time in novel zone, NPP test1.61.73.71.11.9-0.3
Total zone transitions, NPP test1.42.03.6-1.6-1.90.1
Total locomotion distance, NPP test0.91.21.9-1.3-1.4-0.2
Locomotion velocity, NPP test1.21.72.7-1.5-1.6-0.4
Intraocular pressure1.01.42.51.61.2-0.5
Delta distance traveled before/after self-admin0.30.40.60.80.70.1
Delta ambulatory time before/after self-admin0.40.50.80.90.80.3
Parametrial fat weight6.014.020.9-4.6-4.21.9
Pavlov. Cond. lever latency0.91.11.71.31.30.2
Pavlov. Cond. magazine entry latency1.62.02.2-1.5-1.3-1.5
Pavlov. Cond. change in total contacts0.50.50.8-0.6-0.7-0.9
Pavlov. Cond. index score0.70.91.2-1.1-1.1-0.5
Pavlov. Cond. latency score1.21.52.2-1.5-1.4-0.5
Pavlov. Cond. lever contacts1.21.42.1-1.5-1.4-0.6
Pavlov. Cond. magazine entry number0.70.91.60.80.61.3
Pavlov. Cond. intertrial magazine entries0.80.91.50.70.91.2
Pavlov. Cond. lever-magazine prob. diff.0.70.91.3-1.1-1.1-0.5
Pavlov. Cond. response bias0.60.71.1-0.8-0.6-1.1
Pavlov. Cond. lever latency1.01.31.81.01.4-1.1
Pavlov. Cond. magazine entry latency0.91.11.61.31.1-0.6
Pavlov. Cond. change in total contacts0.10.20.5-0.2-0.20.7
Pavlov. Cond. index score0.10.10.30.1-0.00.6
Pavlov. Cond. latency score0.10.10.2-0.1-0.20.4
Pavlov. Cond. lever contacts0.91.32.0-1.0-1.40.9
Pavlov. Cond. magazine entry number1.61.93.3-1.8-1.5-0.5
Pavlov. Cond. intertrial magazine entries2.84.26.6-2.6-2.4-0.5
Pavlov. Cond. lever-magazine prob. diff.0.10.10.10.30.20.3
Pavlov. Cond. response bias0.40.51.40.5-0.11.2
Time in closed arm before self-admin0.40.50.60.70.80.5
Time in closed arm after self-admin2.62.84.62.21.90.5
Time in open arm before self-admin0.30.41.10.1-0.0-1.0
Time in open arm after self-admin2.93.25.1-2.3-2.10.2
Bone: periosteal estimation3.47.19.8-3.1-3.01.6
Bone: periosteal perimeter1.52.74.3-2.1-1.9-0.1
Bone: post-yield work4.26.210.0-2.9-3.20.4
Progressive ratio test 1 active lever presses0.10.10.30.5-0.0-0.2
Progressive ratio test 1 breakpoint0.10.10.40.60.0-0.2
Progressive ratio test 1 inactive lever presses0.00.00.0-0.1-0.2-0.2
Progressive ratio test 2 active lever presses1.31.53.21.81.1-0.1
Progressive ratio test 2 breakpoint1.11.32.81.71.0-0.1
Progressive ratio test 2 inactive lever presses0.10.20.30.30.2-0.6
Incentive Sensitization - Responses0.50.70.80.90.9-0.8
Incentive Sensitization - Breakpoint0.40.50.60.70.6-0.8
Sum of active levers in priming session hrs 5-61.71.62.0-1.1-1.2-1.4
Progressive ratio0.60.92.1-0.6-0.5-1.4
Food seeking constrained by brief footshock0.10.20.5-0.10.1-0.7
Seeking ratio, punishment vs. effort0.00.00.1-0.1-0.00.3
Retroperitoneal fat weight1.46.08.8-2.9-3.0-0.8
Reaction time number correct0.91.63.01.71.3-0.4
Reaction time mean minus median0.00.10.2-0.3-0.4-0.1
Reaction time mean minus median AUC0.00.10.10.40.2-0.1
Reaction time num false alarms0.20.30.50.30.70.4
Reaction time num false alarms AUC0.00.00.0-0.10.20.1
Reaction time trials correct on left0.91.63.01.71.3-0.4
Reaction time trials on left0.81.62.91.71.3-0.4
Reaction time mean0.71.13.1-0.2-0.3-1.7
Reaction time mean AUC0.71.23.50.0-0.1-1.9
Median of all reaction times1.01.44.1-0.1-0.2-2.0
Reaction time omissions0.60.82.2-0.1-0.5-1.5
Reaction time false alarm rate0.20.20.5-0.7-0.10.4
Reaction time premature initiation rate0.71.03.00.30.1-1.7
Reaction time premature initiations1.12.04.01.20.8-2.0
Std. dev. reaction times0.20.30.7-0.4-0.4-0.8
Reaction time trials completed0.81.62.91.71.3-0.4
Reaction time trials AUC0.71.32.31.51.2-0.4
Run reversals in cocaine runway, females1.81.82.8-1.7-1.4-0.9
Run reversals in cocaine runway, males0.10.10.20.40.50.2
Latency to leave start box in cocaine runway1.21.42.6-0.8-1.01.6
Latency to leave start box in cocaine runway, F0.70.82.20.60.01.5
Latency to leave start box in cocaine runway, M1.81.72.8-1.7-1.30.7
Std. dev. time between licks in bursts0.10.10.20.4-0.0-0.5
Total sessions with >9 infusions0.30.40.9-0.4-0.9-0.4
Short access day 1 total inactive lever presses0.50.71.90.0-0.2-1.4
Short access day 10 total inactive lever presses0.41.01.91.41.00.5
Short access day 1 total infusions1.31.82.91.30.9-1.7
Short access day 10 total infusions0.10.20.40.7-0.10.1
Short access total infusions0.20.20.40.2-0.5-0.6
Short access day 1 locomotion1.31.63.31.81.20.1
Short access day 10 total locomotion0.40.72.1-0.3-0.31.4
Short access total locomotion0.50.60.8-0.4-0.90.9
One hour access (0.3 mA shock)0.10.10.2-0.4-0.2-0.0
Compulsive drug intake1.21.52.9-1.7-1.0-0.7
One hour access (shock baseline)0.20.40.70.80.30.6
Social response proportion0.50.61.00.61.0-0.7
Social responses0.20.30.40.50.6-0.6
Social time0.70.91.10.90.9-1.0
Soleus weight3.610.515.9-4.0-3.81.1
Bone: stiffness4.98.010.9-3.3-3.02.0
Tibialis anterior weight6.722.637.1-5.5-6.1-0.3
Time to tail flick, vehicle, before self-admin1.11.21.9-1.4-1.2-0.4
Time to tail flick, vehicle, after self-admin0.10.10.2-0.4-0.1-0.3
Time to tail flick, test, before self-admin1.41.52.7-1.1-1.7-0.6
Time to tail flick, test, after self-admin0.80.92.30.40.31.5
Delta time to tail flick, vehicle, before/after SA0.10.10.3-0.1-0.10.5
Delta time to tail flick, test, before/after SA0.60.71.50.30.61.2
Tibia length0.10.20.4-0.6-0.50.0
Ambulatory time before self-admin0.71.11.71.21.30.5
Ambulatory time after self-admin1.72.03.11.71.8-0.2
Bone: tissue strength0.20.20.4-0.2-0.3-0.6
Total heroin consumption0.80.92.1-0.7-0.4-1.4
Bone: trabecular number0.20.40.5-0.5-0.70.5
Bone: trabecular spacing0.71.02.60.40.5-1.6
Bone: trabecular thickness1.21.42.0-1.4-1.20.9
Bone: trabecular tissue density0.81.11.9-1.4-1.0-0.7
Water consumed over 24 hours1.01.32.21.51.10.6
Number of responses in last shaping day1.11.22.90.80.2-1.7