1. Neuroscience

Newly trained navigation and verbal memory skills in humans elicit changes in task-related networks but not brain structure

  1. Li Zheng
  2. Zachary Boogaart
  3. Andrew McAvan
  4. Joshua Garren
  5. Stephanie G Doner
  6. Bradley J Wilkes
  7. Will Groves
  8. Ece Yuksel
  9. Lucia Cherep
  10. Arne Ekstrom  Is a corresponding author
  11. Steven M Weisberg  Is a corresponding author
  1. Department of Psychology, University of Arizona, United States
  2. Evelyn McKnight Brain Institute, University of Arizona, United States
  3. The University of Miami Miller School of Medicine, University of Miami, United States
  4. Department of Applied Physiology & Kinesiology, University of Florida, United States
  5. Department of Psychology, University of Florida, United States
  6. Evelyn McKnight Brain Institute, University of Florida, United States
https://doi.org/10.7554/eLife.106873.3
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Cite this article
  1. Li Zheng
  2. Zachary Boogaart
  3. Andrew McAvan
  4. Joshua Garren
  5. Stephanie G Doner
  6. Bradley J Wilkes
  7. Will Groves
  8. Ece Yuksel
  9. Lucia Cherep
  10. Arne Ekstrom
  11. Steven M Weisberg
(2025)
Newly trained navigation and verbal memory skills in humans elicit changes in task-related networks but not brain structure
eLife 14:RP106873.
https://doi.org/10.7554/eLife.106873.3
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14 figures, 17 tables and 1 additional file

Figures

Figure 1
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Experimental design.

Participants were randomly assigned to one of three training conditions as follows. (a) In the verbal memory training condition (n=27), participants underwent 10 sessions of verbal memory training involving word free recall after a distractor task, with list complexity increasing over sessions. To quantify the training effect, we conducted a linear regression analysis on the maximum number of correctly recalled words for each day. The slopes of the regression lines for the first 5 days and the last 5 days were calculated separately, and both slopes were significantly greater than zero, indicating a significant improvement in memory performance over the training period: first 5 days, t(26) = 16.971, p<0.001; last 5 days, t(26) = 23.579, p<0.001. (b) In the navigation training condition (n=27), participants trained in a large virtual environment, navigating between buildings until optimal paths were learned. Subsections of the environment were integrated progressively across sessions. Participants improved over the course of the training, showing higher error initially for subsection A (M=79.09, SD = 33.22) compared to subsection B (M=65.52, SD = 26.89), which was significantly reduced for subsection C (M=47.44, SD = 31.22). (c) In the active control condition (n=21), participants watched videos related to memory and navigation, answering multiple-choice questions afterward, with accuracy consistently above 50%, indicating engagement. Training schedules spanned 4–8 weeks, with each session lasting 2 hr. Notes: Boxplots are centered on the median, boxes extend to first and third quartiles, whiskers extend to 1.5 times the interquartile range or minima/maxima in the absence of outliers. Each individual dot represents data from an individual subject. Red diamonds represent the mean value.

Figure 2
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Changes in learning rates across training conditions from pre-test to post-test.

(a) Learning rate for spatial navigation improvement on the Navigation Transfer task. All three groups improved from pre-test to post-test, with the Navigation group demonstrating the largest effect (paired-sample t-tests; Navigation: t(26) = 4.43, p<0.001, Cohen’s d=1.32; Verbal Memory: t(26) = 2.31, p=0.03, Cohen’s d=0.596; Video Control: t(18) = 3.97, p<0.001, Cohen’s d=1.20.) (b) Learning rate for verbal memory improvement in the Verbal Memory transfer task. Only the Verbal Memory group significantly improved from pre-test to post-test (paired-sample t-tests; Verbal Memory: t(25) = 3.32, p=0.003, Cohen’s d=0.608; Navigation: t(26) = 0.488, p=0.63, Cohen’s d=0.113; Video Control: t(20) = –0.55, p=0.588, Cohen’s d=0.164). Each individual dot represents data from an individual subject. Red diamonds represent the mean value. *p<0.05, **p<0.01, ***p<0.001.

Figure 3
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Relationship between medial temporal lobe volumes and training-induced learning rate changes across conditions.

(a) Normalized MTL subregion volumes across conditions (Verbal Memory/Navigation/Video Control) and sessions (pre-test/post-test). No significant changes were detected in CA1, CA23DG, SUB, ERC, PRC, or PHC volumes between pre-test and post-test (paired-sample t-tests, all ps >0.588, FDR-corrected; Verbal Memory: n=26, Navigation: n=27, Video Control: n=21). (b) Normalized hippocampal volumes (HIP), including left and right hippocampus, across conditions and sessions. No significant changes in total, left, or right hippocampal volume from pre-test to post-test (paired-sample t-tests, all ps >0.256, FDR-corrected; Verbal Memory: n=26, Navigation: n=27, Video Control: n=21). (c) Correlations between changes in learning rate (post-test minus pre-test) and average CA23DG volume across groups. A significant positive correlation was observed in the Verbal Memory group (partial correlation controlling for sex and site: r(23) = 0.493, p=0.017, FDR corrected), suggesting that participants with larger CA23DG volumes exhibited greater improvements in verbal memory performance. No such correlation was observed in the Navigation group (r(25) = –0.083, p=0.953, FDR corrected) or Video Control group (r(19) = –0.12, p=0.953, FDR corrected). (d) Correlations between the changes in learning rate (post-test minus pre-test) and average total hippocampal volume across groups. A significant positive correlation was observed only in the Verbal Memory group (partial correlation controlling for sex and site: r(23) = 0.605, p=0.006, FDR corrected). No such correlation was observed in the Navigation group (r(25) = 0.038, p=0.858, FDR corrected) or Video Control group (r(19) = –0.123, p=0.858, FDR corrected). Boxplots are centered on the median, boxes extend to first and third quartiles, whiskers extend to 1.5 times the interquartile range or minima/maxima in the absence of outliers. Each individual dot represents data from an individual subject. Red diamonds represent the mean value.

Figure 4
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Task-related informational connectivity changes during encoding as a result of verbal memory and navigation interventions.

(a) Representational similarity matrices (RSMs) that illustrate within-context correlations (spatial and temporal) for each of the 242 brain ROIs. Informational connectivity between 242 ROIs was derived by correlating the RSMs across regions within different contexts, resulting in three types of informational connectivity matrices (ICMs): the spatial ICM, the temporal ICM, and the combined ICM for all within-context trials. (b) Visualization of the 242 predefined ROIs, color-coded by functional networks. (c) Schematic of the experimental design for the source memory task during the encoding stage, highlighting tasks related to spatial and temporal contexts. (d) Differences in task-related informational connectivity during encoding (Verbal Memory (n=25)>Navigation (n=26)+Video (n=20); Navigation >Verbal Memory + Video) across training conditions. Results are shown for all trials, as well as separately for spatial and temporal encoding contexts. Red lines indicate regions with significantly increased connectivity (post >pre), while blue lines indicate regions with significantly decreased connectivity (post <pre). The top panels display results for the Verbal Memory group, while the bottom panels display results for the Navigation group. All Pearson correlation coefficients (r values) were Fisher-Z transformed prior to statistical analysis. Significance thresholds for informational connectivity were determined using 10,000 permutation simulations. The reported results have been corrected for multiple comparisons using the FDR method, with a q-value threshold of less than 0.05. Nav: Navigation.

Figure 5
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Task-related informational connectivity changes during source retrieval as a result of verbal memory and navigation interventions.

(a) Schematic of the source memory task structure during retrieval. Participants were scanned during source retrieval, in which they identified whether an item was previously seen and determined its spatial or temporal context. (b) Differences in task-related informational connectivity during source retrieval (Verbal Memory (n=24)>Navigation (n=22)+Video (n=20); Navigation >Verbal Memory + Video) across training conditions. Results are shown for all trials, as well as separately for spatial and temporal encoding contexts. Red lines indicate regions with significantly increased connectivity (post >pre), while blue lines indicate regions with significantly decreased connectivity (post <pre). The top panels display results for the Verbal Memory group, while the bottom panels display results for the Navigation group. All Pearson correlation coefficients (r values) were Fisher-Z transformed prior to statistical analysis. Significance thresholds for informational connectivity were determined using 10,000 permutation simulations. The reported results have been corrected for multiple comparisons using the FDR method, with a q-value threshold of less than 0.05. Nav: Navigation.

Figure 6
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Multivariate informational connectivity pattern distance between pre-test and post-test.

(a) Schematic of multivariate distance analysis. For each ROI, informational connectivity between the current ROI and the remaining 214 ROIs was extracted from both spatial and temporal ICMs, and distances were calculated as 1 minus the Pearson or Spearman correlation coefficient. (b) and (c) Among all 242 ROIs, only the SFG showed a greater distance between spatial and temporal ICMs in post-test compared to pre-test in the Verbal Memory (n=24) group, but not in the Navigation (n=22) or Video Control (n=20) groups. ICM: informational connectivity matrix. Paired t-tests were conducted to evaluate statistical differences between pre and post. *p<0.05, FDR corrected.

Appendix 3—figure 1
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Schematic representations of additional formats of the Navigation Transfer Task.

(a) Navigation Pointing Task. (b) Navigation Model Building Task.

Appendix 3—figure 2
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Performance on the Navigation Pointing task and Navigation Model Building task.

(a) Paired-sample t-tests indicated that all three groups demonstrated a decrease in overall pointing error from pre-test to post-test, with the Verbal Memory group showing the largest effect (Navigation: t(26) = 4.76, p<0.001, Cohen’s d=0.925; Verbal Memory: t(24) = 5.97, p<0.001, Cohen’s d=0.983; Video Control: t(20) = 5.55, p<0.001, Cohen’s d=0.758). (b) Paired-sample t-tests indicated that all three groups demonstrated a decrease in between-environment pointing error from pre to post-test (Navigation: t(26) = 5.16, p<0.001, Cohen’s d=1.07; Verbal Memory: t(24) = 5.88, p<0.001, Cohen’s d=1.13; Video Control: t(20) = 4.22, p<0.001, Cohen’s d=0.871). (c) Paired-sample t-tests indicated that all three groups demonstrated a decrease in within-environment pointing error from pre to post (Navigation: t(26) = 2.36, p=0.03, Cohen’s d=0.479; Verbal Memory: t(24) = 3.74, p=0.001, Cohen’s d=0.5; Video Control: t(20) = 3.34, p=0.003, Cohen’s d=0.435). (d) Paired-sample t-tests indicated that all three groups demonstrated an improvement in map accuracy from pre to post-test (Navigation: t(26) = 4.95, p<0.001, Cohen’s d=0.81; Verbal Memory: t(26) = 5.72, p<0.001, Cohen’s d=0.94; Video Control: t(20) = 2.97, p=0.008, Cohen’s d=0.66). Each individual dot represents data from an individual subject. Red diamonds represent the mean value.

Appendix 3—figure 3
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Left and right hippocampal volumes were correlated with changes in Verbal Memory Transfer task performance between pre-test and post-test.

(a) A significant positive correlation was observed between the learning rate in the Verbal Memory group and right hippocampal volume (r(23) = 0.601, p-FDR=0.007), while no significant correlations were found for the Navigation (r(25) = 0.008, p-FDR=0.970) or Video Control group (r(19) = –0.163, p=0.758), after controlling for sex and site as covariates. (b) A significant positive correlation was observed between the learning rate in the Verbal Memory group and left hippocampal volume (r(23) = 0.568, p-FDR=0.014), while no significant correlations were found for the Navigation (r(25) = 0.038, p-FDR=0.858) or Video Control group (left: r(19) = –0.123, p-FDR=0.858), after controlling for sex and site as covariates.

Appendix 3—figure 4
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Memory performance (hit rate and false alarm rate) during the Source Memory task in the scanner.

(a) Paired-sample t-tests indicated that none of the three conditions demonstrated a significant increase in hit rate from pre to post-test: Verbal Memory: t(23) = 1.429, p=0.166, Cohen’s d=0.223; Navigation: t(21) = 1.847, p=0.079, Cohen’s d=0.316; Control: t(19) = 1.082, p=0.293, Cohen’s d=0.208. (b) Paired-sample t-tests showed no significant changes in FA across sessions for any condition: Verbal Memory: t(23) = 1.477, p=0.153, Cohen’s d=0.451; Navigation: t(21) = –0.699, p=0.492, Cohen’s d=–0.168; Control: t(19) = –0.0005, p=0.999, Cohen’s d=–0.0001. (c) Paired-sample t-tests indicated no significant increase in hit rate from pre-test to post-test in any condition: Verbal Memory: t(23) = 1.102, p=0.282, Cohen’s d=0.217; Navigation: t(21) = 1.690, p=0.106, Cohen’s d=0.304; Control: t(19) = 1.433, p=0.168, Cohen’s d=0.214. (d) Paired-sample t-tests revealed no significant changes in hit rate across sessions: Verbal Memory: t(23) = 1.288, p=0.210, Cohen’s d=0.196; Navigation: t(21) = 1.513, p=0.145, Cohen’s d=0.288; Control: t(19) = 0.564, p=0.580, Cohen’s d=0.147.

Appendix 3—figure 5
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Memory performance (reaction time) during the Source Memory task in the scanner.

(a) Paired-sample t-tests showed no significant changes in RT from pre-test to post-test for the Navigation condition (t(21) = –0.453, p=0.655, Cohen’s d=–0.107) or the Video Control condition (t(19) = 1.454, p=0.162, Cohen’s d=0.363). However, the Verbal Memory condition demonstrated a marginally significant decrease in RT from pre to post-test (t(23) = 2.042, p=0.053, uncorrected, Cohen’s d=0.447). (b) For spatial source memory, paired-sample t-tests showed no statistically significant RT changes across conditions: Verbal Memory (t(23) = 1.444, p=0.162, Cohen’s d=0.321), Navigation (t(21) = –0.189, p=0.852, Cohen’s d=–0.041), and Video Control (t(19) = 1.073, p=0.297, Cohen’s d=0.264). (c) For temporal source memory, paired-sample t-tests showed that the Verbal Memory group demonstrated a trend toward decreased RT in temporal source memory from pre to post-test (t(23) = 1.910, p=0.069, uncorrected, Cohen’s d=0.434), while no such trends were observed in the Navigation (t(21) = –0.622, p=0.541, Cohen’s d=–0.163) or Video Control conditions (t(19) = 1.708, p=0.103, Cohen’s d=0.445).

Appendix 3—figure 6
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Univariate activation changes from pre-test to post-test during Source Memory task encoding.

(a) In the Verbal Memory condition (n=25), a significant decrease in activation was observed in several regions during the post-test compared to the pre-test (post <pre). These regions included the left dorsolateral occipital cortex (dLOC; Z=4.66, MNI: −46,–62, 30), the left middle frontal gyrus (MFG; Z=4.32, MNI: –42, 12, 50), the left frontal pole (Z=4.20, MNI: –24, 40, 46), the left middle temporal gyrus (MTG; Z=4.28, MNI: −60,–24, –12), and the left precuneus (Z=3.80, MNI: −10,–52, 38). (b) In the Navigation condition (n=26), post-test activation was significantly increased in the right middle temporal gyrus (MTG; Z=4.03, MNI: 42,–54, 6) compared to the pre-test (post >pre). (c) In the Verbal Memory condition (n=25), a significant decrease in spatial encoding activation was observed during the post-test compared to the pre-test (LOC; Z=3.97, MNI: −44,–62, 32) (d) In the Verbal Memory group (n=25), a decrease in temporal encoding activation was observed in the left LOC (Z=3.97, MNI: −40,–62, 32) during the post-training stage relative to the pre-test. All results were derived from whole-brain voxel-wise paired t-tests, and statistical maps were thresholded using cluster-based detection methods with a voxel-wise threshold of z>3.1 and a cluster-level significance threshold of p<0.05, corrected for multiple comparisons across the whole brain using Gaussian Random Field Theory.

Appendix 3—figure 7
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Univariate activation during the encoding stage of the Source Memory task, presented separately for pre-test and post-test sessions.

Whole-brain mixed-effects analyses were conducted, and group-level statistical maps were thresholded using cluster-based detection methods with a voxel-wise height threshold of z>3.1 and a cluster-level significance threshold of p<0.05, corrected for multiple comparisons across the whole brain using Gaussian Random Field Theory. Results are shown for the Verbal Memory (n=25), Navigation (n=26), and Video Control (n=20) groups.

Appendix 3—figure 8
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Univariate activation during the retrieval stage of the Source Memory task, presented separately for pre-test and post-test sessions.

Whole-brain mixed-effects analyses were conducted, and group-level statistical maps were thresholded using cluster-based detection methods with a voxel-wise height threshold of z>3.1 and a cluster-level significance threshold of p<0.05, corrected for multiple comparisons across the whole brain using Gaussian Random Field Theory. Results are shown for the Verbal Memory (n=24), Navigation (n=22), and Video Control (n=20) groups.

Tables

Key resources table
Reagent type (species) or resourceDesignationSource or referenceIdentifiersAdditional information
Software, algorithmPsychoPy v2022.2https://www.psychopy.orgRRID:SCR_006571Used for behavioral task presentation
Software, algorithmUnity 3D (2021)https://unity.comUsed to develop the virtual navigation environment
Software, algorithmFSL v5.0.1https://fsl.fmrib.ox.ac.ukRRID:SCR_002823Used for preprocessing and registration of MRI data
Software, algorithmAdvanced Normalization Toolshttps://stnava.github.io/ANTs/RRID:SCR_004757Used for registration fMRI and DWI data
Software, algorithmMRtrix3https://www.mrtrix.org/RRID:SCR_024123Used for DWI data analysis
Software, algorithmAutomatic hippocampal subfield segmentationhttps://sites.google.com/view/ashs-dox/homeRRID:SCR_005996Used for hippocampus segmentation
Software, algorithmFreeSurfer v7.4.1https://surfer.nmr.mgh.harvard.eduRRID:SCR_001847Used for cortical reconstruction and volumetric segmentation
Software, algorithmMATLABhttps://www.mathworks.com/products/matlab.htmlRRID:SCR_001622Used for custom scripts for behavioral and fMRI data analysis
Software, algorithmRhttps://www.r-project.org/RRID:SCR_001905Used for custom scripts for figures and statistical analysis
Appendix 2—table 1
Demographics information.
Verbal MemoryNavigationVideo ControlAll
Sample size27272175
SexMale711725
Female20161450
SiteSite 113131339
Site 21414836
Male-Site 124511
Female-Site 199826
Male-Site 257214
Female-Site 297622
AgeAge (min-max/Year)20.89 (18-26)22.11 (18-32)22 (18–32)21.67
Age-Site120.3123.0023.0022.10
Age-Site221.4321.2920.3821.03
Appendix 2—table 2
Sample size information by condition and test, reflecting exclusions due to outlier performance, excessive head movement during scanning, or missing data.
TestVerbal MemoryNavigationVideo
Verbal Memory Training272721
Navigation Training272721
Verbal Memory Transfer Task262721
Navigation Transfer Task272719
Navigation Pointing Task252721
Navigation Model Building Task272721
Source Memory Task-Encoding252620
Source Memory Task-Retrieval242220
Hippocampal volume262721
DWI202218
Appendix 2—table 3
Study Timeline.
Pre-test(Day 1)Training(Days 2–11)Post-test(Day 12)
Complete consent, MRI prescreen, and demographics surveyComprehensive training varies based on which condition the participant was assignedReview consent, MRI prescreen
Navigation Transfer task (Virtual Silcton, Unity)Navigation Task (navigation training in virtual Arida, Unity)Navigation Transfer task (Virtual Silcton, Unity)
Navigation Pointing task (Virtual Silcton, web-based)Verbal memory Task (verbal memory training with the modified method of loci, Unity)Navigation Pointing task (Virtual Silcton, web-based)
Navigation Model Building task (Virtual Silcton, web-based)Control task (video control by viewing informative videos and answering questions, Qualtrics)Navigation Model Building task (Virtual Silcton, web-based)
Verbal Memory Transfer task (Psychopy)Verbal Memory Transfer task (PsychoPy)
Attention task (web-based)Attention task (web-based)
Source Memory task (PsychoPy, fMRI scanned)Source Memory task (PsychoPy, fMRI scanned)
Debrief survey (Quatrics)
Appendix 2—table 4
Correlations between the volumes of MTL subregions and changes in learning rate from pre-test to post-test in the Verbal Memory Transfer task.
ConditionROIrpMethod
Verbal Memory
N = 25		Ant-HIP0.2480.232Pearson
Post-HIP–0.0430.837Spearman
CA10.1380.51Pearson
CA23DG0.5320.006*Pearson
SUB0.0250.907Pearson
ERC0.0690.743Pearson
PRC0.1620.438Pearson
PHC–0.0680.748Spearman
Navigation
N = 27		Ant-HIP–0.0640.751Spearman
Post-HIP0.1830.362Pearson
CA10.0840.676Pearson
CA23DG–0.0120.953Pearson
SUB0.0350.863Pearson
ERC–0.1420.48Pearson
PRC0.0230.91Pearson
PHC0.2230.264Pearson
Video
N = 21		Ant-HIP–0.1050.651Pearson
Post-HIP0.0350.88Pearson
CA1–0.1310.572Pearson
CA23DG–0.0830.719Pearson
SUB–0.240.294Pearson
ERC0.1490.52Spearman
PRC0.1210.6Pearson
PHC–0.2920.198Pearson
  1. *

    Significant results after FDR correction. Ant: anterior; Post: posterior

Appendix 2—table 5
Correlations between the volumes of MTL subregions and changes in learning rate from pre-test to post-test in the Navigation Transfer task.
ConditionROIrpMethod
Verbal Memory
N = 26		Ant-HIP–0.3280.102Pearson
Post-HIP–0.1470.471Spearman
HIP–0.3140.119Pearson
LHIP–0.2910.149Pearson
RHIP–0.3050.129Pearson
CA1–0.1810.377Pearson
CA23DG–0.2610.198Pearson
SUB–0.2570.205Pearson
ERC0.0270.897Pearson
PRC–0.0180.931Pearson
PHC–0.2430.23Spearman
Navigation
N = 27		Ant-HIP–0.4820.012Spearman
Post-HIP–0.0420.837Pearson
HIP–0.2660.181Pearson
LHIP–0.2470.214Pearson
RHIP–0.2740.167Pearson
CA1–0.2460.215Pearson
CA23DG–0.3340.088Pearson
SUB0.0870.665Pearson
ERC0.0080.967Pearson
PRC–0.0210.917Pearson
PHC0.0280.891Pearson
Video
N = 19		Ant-HIP–0.1010.681Pearson
Post-HIP0.0480.846Pearson
HIP0.1240.612Pearson
LHIP0.2460.31Pearson
RHIP0.0280.911Spearman
CA10.020.934Pearson
CA23DG0.1280.6Pearson
SUB0.0530.828Pearson
ERC0.1940.425Pearson
PRC0.240.322Pearson
PHC–0.0890.716Pearson

  1. Ant: anterior; Post: posterior

Appendix 2—table 6
Correlations between the volumes of MTL subregions and changes in the average number of correctly recalled words from pre-test to post-test in the Verbal Memory Transfer task.
ConditionROIrpMethod
Verbal Memory
N = 25		Ant-HIP–0.3500.087Pearson
Post-HIP–0.0420.842Spearman
HIP–0.2350.258Pearson
LHIP–0.1720.410Pearson
RHIP–0.2670.197Pearson
CA1–0.2120.309Pearson
CA23DG–0.3290.109Pearson
SUB0.1060.615Pearson
ERC0.0300.888Pearson
PRC–0.2190.292Pearson
PHC–0.2020.334Spearman
Navigation
N = 27		Ant-HIP0.1220.545Spearman
Post-HIP–0.1310.516Pearson
HIP0.0120.953Pearson
LHIP–0.0320.875Pearson
RHIP0.0490.809Pearson
CA10.0230.911Pearson
CA23DG0.0430.830Pearson
SUB–0.0850.674Pearson
ERC0.1350.503Pearson
PRC–0.0620.757Pearson
PHC–0.2560.197Pearson
Video
N = 21		Ant-HIP–0.1110.633Pearson
Post-HIP–0.0120.959Pearson
HIP–0.1250.589Pearson
LHIP–0.1560.499Pearson
RHIP–0.1830.426Spearman
CA10.1160.616Pearson
CA23DG–0.1470.525Pearson
SUB–0.2120.357Pearson
ERC–0.3650.104Spearman
PRC–0.1890.413Pearson
PHC–0.0880.703Pearson

  1. Ant: anterior; Post: posterior

Appendix 2—table 7
Correlations between the volumes of MTL subregions and changes in the number of trials to criterion from pre-test to post-test in the Verbal Memory Transfer task.
ConditionROIrpMethod
Verbal Memory
N = 25		Ant-HIP0.0950.653Spearman
Post-HIP0.3160.124Spearman
HIP–0.1630.438Spearman
LHIP–0.1110.598Spearman
RHIP–0.2230.285Spearman
CA10.0280.896Spearman
CA23DG0.0740.726Spearman
SUB–0.3980.049Spearman
ERC–0.3340.103Spearman
PRC0.1390.509Spearman
PHC0.0410.846Spearman
Navigation
N = 27		Ant-HIP0.0840.679Spearman
Post-HIP0.0080.970Spearman
HIP–0.0230.908Spearman
LHIP–0.0420.837Spearman
RHIP–0.0330.872Spearman
CA1–0.0150.943Spearman
CA23DG–0.0280.890Spearman
SUB0.0990.622Spearman
ERC–0.0110.956Spearman
PRC0.1370.495Spearman
PHC0.2570.196Spearman
Video
N = 21		Ant-HIP0.2230.332Pearson
Post-HIP0.2420.290Pearson
HIP0.3590.110Pearson
LHIP0.4130.063Pearson
RHIP0.2220.334Spearman
CA10.1670.471Pearson
CA23DG0.1780.440Pearson
SUB0.4360.048Pearson
ERC0.2020.379Spearman
PRC0.0720.756Pearson
PHC0.4610.035Pearson

  1. Ant: anterior; Post: posterior

Appendix 2—table 8
Correlations between the volumes of MTL subregions and changes in slope from pre-test to post-test in the Verbal Memory Transfer task.
ConditionROIrpMethod
Verbal Memory
N = 26		Ant-HIP0.300.15Pearson
Post-HIP–0.140.51Spearman
HIP0.390.05Pearson
LHIP0.320.12Pearson
RHIP0.420.04Pearson
CA10.130.54Pearson
CA23DG0.500.01Pearson
SUB0.150.46Pearson
ERC0.110.59Pearson
PRC0.190.37Pearson
PHC0.010.95Spearman
Navigation
N = 27		Ant-HIP0.040.83Spearman
Post-HIP0.230.26Pearson
HIP0.090.67Pearson
LHIP0.100.63Pearson
RHIP0.070.71Pearson
CA10.180.36Pearson
CA23DG0.020.92Pearson
SUB0.060.75Pearson
ERC–0.100.63Pearson
PRC0.010.97Pearson
PHC0.210.30Pearson
Video
N = 21		Ant-HIP–0.230.32Pearson
Post-HIP–0.050.82Pearson
HIP–0.290.20Pearson
LHIP–0.280.21Pearson
RHIP–0.280.21Spearman
CA1–0.160.50Pearson
CA23DG–0.150.51Pearson
SUB–0.310.17Pearson
ERC0.090.71Spearman
PRC0.050.82Pearson
PHC–0.330.14Pearson

  1. Ant: anterior; Post: posterior

Appendix 2—table 9
Correlations between the volumes of MTL subregions and changes in path errors from pre-test to post-test in the Navigation Transfer task.
ConditionROIrpMethod
Verbal Memory
N = 26		Ant-HIP0.0800.698Pearson
Post-HIP0.1170.570Spearman
HIP0.0410.844Pearson
LHIP0.1170.570Pearson
RHIP0.0410.844Pearson
CA10.1630.426Pearson
CA23DG–0.2140.294Pearson
ERC0.2740.176Pearson
PHC0.0470.820Pearson
PRC–0.4520.020Pearson
SUB0.3960.045Spearman
Navigation
N = 27		Ant-HIP–0.1370.493Spearman
Post-HIP–0.1810.366Pearson
HIP–0.0580.775Pearson
LHIP–0.0320.873Pearson
RHIP–0.0780.701Pearson
CA1–0.2940.137Pearson
CA23DG0.0110.956Pearson
ERC–0.0720.723Pearson
PHC0.0270.892Pearson
PRC–0.0830.681Pearson
SUB0.1010.617Pearson
Video
N = 19		Ant-HIP0.3010.211Pearson
Post-HIP0.2100.389Pearson
HIP–0.1930.429Pearson
LHIP–0.1870.443Pearson
RHIP–0.1300.595Spearman
CA1–0.1860.446Pearson
CA23DG0.0520.833Pearson
ERC0.0650.791Pearson
PHC–0.2910.227Pearson
PRC0.1500.540Pearson
SUB–0.2770.252Pearson

  1. Ant: anterior; Post: posterior

Appendix 2—table 10
Correlations between the volumes of MTL subregions and changes in overall pointing errors from pre-test to post-test in the Navigation Pointing Error task.
ConditionROIrpMethod
Verbal Memory
N=24		Ant-HIP–0.1430.504Pearson
Post-HIP0.1910.369Spearman
HIP–0.2050.336Pearson
LHIP–0.1270.554Pearson
RHIP–0.2520.236Pearson
CA10.0001.000Pearson
CA23DG–0.2010.346Pearson
ERC–0.0860.690Pearson
PHC–0.1340.531Spearman
PRC0.3140.136Pearson
SUB–0.2630.214Pearson
Navigation
N=27		Ant-HIP–0.1190.553Spearman
Post-HIP–0.3170.107Pearson
HIP–0.1590.429Pearson
LHIP–0.1980.322Pearson
RHIP–0.1210.548Pearson
CA1–0.2950.136Pearson
CA23DG–0.0170.932Pearson
ERC–0.0240.907Pearson
PHC–0.2270.256Pearson
PRC–0.0900.656Pearson
SUB–0.2080.299Pearson
Video
N=21		Ant-HIP–0.1550.503Pearson
Post-HIP0.0410.860Pearson
HIP0.2000.385Pearson
LHIP0.2470.280Pearson
RHIP0.0420.859Spearman
CA1–0.0400.864Pearson
CA23DG0.1880.415Pearson
ERC–0.2460.282Spearman
PHC0.2470.282Pearson
PRC0.2380.300Pearson
SUB0.2500.274Pearson
Appendix 2—table 11
Correlations between the volumes of MTL subregions and changes in within-environment pointing error from pre-test to post-test in the Navigation Pointing Error task.
ConditionROIrpMethod
Verbal Memory N=24Ant-HIP–0.1420.508Pearson
Ant-HIP0.1650.439Spearman
HIP0.1100.610Pearson
LHIP0.1630.447Pearson
RHIP0.0550.798Pearson
CA10.0400.854Pearson
CA23DG0.2390.261Pearson
ERC0.1920.369Pearson
PHC–0.1450.497Spearman
PRC0.4110.046Pearson
SUB–0.1490.488Pearson
Navigation N=27Ant-HIP–0.2420.222Spearman
Post-HIP–0.2490.211Pearson
HIP–0.1660.408Pearson
LHIP–0.1830.360Pearson
RHIP–0.1470.466Pearson
CA1–0.1990.321Pearson
CA23DG–0.1600.427Pearson
ERC0.0190.925Pearson
PHC–0.1780.374Pearson
PRC–0.2040.309Pearson
SUB–0.0100.961Pearson
Video
N=21		Ant-HIP0.1660.471Pearson
Post-HIP0.1810.432Pearson
HIP0.0600.796Pearson
LHIP0.0400.865Pearson
RHIP0.0440.850Spearman
CA1–0.0780.738Pearson
CA23DG0.3010.185Pearson
ERC–0.2600.254Spearman
PHC–0.1120.629Pearson
PRC0.4150.062Pearson
SUB–0.2320.312Pearson
Appendix 2—table 12
Correlations between the volumes of MTL subregions and changes in between-environment pointing error from pre-test to post-test in the Navigation Pointing Error Task.
ConditionROIrpMethod
Verbal Memory
N=24		Ant-HIP–0.1200.576Pearson
Post-HIP0.1900.373Spearman
HIP–0.2850.177Pearson
LHIP–0.2110.323Pearson
RHIP–0.3210.127Pearson
CA1–0.0140.948Pearson
CA23DG–0.3270.119Pearson
ERC–0.1720.423Pearson
PHC–0.1830.391Spearman
PRC0.2270.286Pearson
SUB–0.2610.218Pearson
Navigation
N=27		Ant-HIP–0.1450.468Spearman
Post-HIP–0.2950.136Pearson
HIP–0.1250.534Pearson
LHIP–0.1690.400Pearson
RHIP–0.0840.676Pearson
CA1–0.2910.141Pearson
CA23DG0.0630.756Pearson
ERC–0.0420.834Pearson
PHC–0.2100.294Pearson
PRC–0.0110.955Pearson
SUB–0.2750.165Pearson
Video
N=21		Ant-HIP–0.2480.278Pearson
Post-HIP–0.0470.838Pearson
HIP0.1820.430Pearson
LHIP0.2430.290Pearson
RHIP–0.0210.930Spearman
CA1–0.0030.989Pearson
CA23DG0.0490.835Pearson
ERC–0.1830.425Spearman
PHC0.3180.160Pearson
PRC0.0440.850Pearson
SUB0.3820.087Pearson
Appendix 2—table 13
Correlations between the volumes of MTL subregions and changes in model-building accuracy from pre-test to post-test in the Navigation Model Building task.
ConditionROIrpMethod
Verbal Memory N=26Ant-HIP0.1250.544Pearson
Post-HIP–0.1580.438Spearman
HIP0.0660.749Pearson
LHIP0.0250.905Pearson
RHIP0.0960.640Pearson
CA1–0.0480.816Pearson
CA23DG0.1370.504Pearson
ERC–0.0090.965Pearson
PHC0.0740.721Spearman
PRC0.3450.085Pearson
SUB0.0170.936Pearson
Navigation
N=27		Ant-HIP–0.0350.863Spearman
Post-HIP0.0080.969Pearson
HIP0.0780.701Pearson
LHIP0.0460.821Pearson
RHIP0.1020.612Pearson
CA10.3220.102Pearson
CA23DG–0.0680.737Pearson
ERC–0.0650.746Pearson
PHC–0.0380.852Pearson
PRC–0.1290.520Pearson
SUB0.0740.715Pearson
Video
N=21		Ant-HIP–0.0830.722Pearson
Post-HIP–0.2310.313Pearson
HIP0.0390.868Pearson
LHIP–0.0200.931Pearson
RHIP0.0920.690Spearman
CA10.1290.577Pearson
CA23DG–0.1810.432Pearson
ERC0.2030.377Spearman
PHC0.0460.842Pearson
PRC–0.4070.067Pearson
SUB0.2350.304Pearson
Appendix 2—table 14
Correlations between the average volumes of MTL subregions and slope from Day 1 to Day 5 in the Verbal Memory Training task.
ConditionROIrpMethod
Verbal Memory N=26Ant-HIP–0.03230.8755Pearson
Post-HIP–0.13210.52Spearman
HIP0.09120.6577Pearson
LHIP0.23910.2395Pearson
RHIP–0.04550.8252Pearson
CA1–0.09930.6292Pearson
CA23DG0.02990.8846Pearson
ERC0.18060.3773Pearson
PHC0.13970.4962Spearman
PRC–0.32570.1044Pearson
SUB0.32390.1065Pearson
Appendix 2—table 15
Correlations between the average volumes of MTL subregions and slope from Day 6 to Day 10 in the Verbal Memory Training task.
ConditionROIrpMethod
Verbal Memory N=26Ant-HIP–0.08820.6682Pearson
Post-HIP–0.05760.78Spearman
HIP–0.11880.5633Pearson
LHIP0.00820.9685Pearson
RHIP–0.21880.2829Pearson
CA1–0.09020.6612Pearson
CA23DG–0.29160.1484Pearson
ERC0.15240.4573Pearson
PHC0.15420.4521Spearman
PRC–0.33930.09Pearson
SUB0.24590.2259Pearson
Appendix 2—table 16
Univariate activation changes from pre-test to post-test during encoding and retrieval, with whole-brain cluster correction and small-volume correction for the hippocampus reported separately.
Pre vs. Post
ConditionContrastResults (Hippocampus)Results (whole brain)
EncodingVideoNoneNone
Verbal MemoryNoneT1>T2: left dLOC (Z=4.66, MNI: −46,–62, 30), left MFG (Z=4.32, MNI: –42, 12, 50), left frontal pole (Z=4.2, MNI: –24, 40, 46), left MTG (Z=4.28, MNI: −60,–24, –12), left precuneus (Z=3.8, MNI: −10,–52, 38).
NavigationNoneT2>T1: right MTG (Z=4.03, MNI: 42,–54, ss6).
Verbal Memory vs. VideoNoneNone
Navigation vs. VideoNoneNone
Verbal Memory vs. NavigationNoneNone
Verbal Memory vs. Video +NavigationNoneNone
Navigation vs. Video +Verbal MemoryNoneNone
Spatial encodingVideoNoneNone
Verbal MemoryNoneT1>T2: Left LOC (Z=3.97, MNI: −44,–62,32), Left frontal pole (Z=3.99, MNI: –18, 48, 36).
NavigationNoneNone
Verbal Memory vs. VideoNoneNone
Navigation vs. VideoNoneNone
Verbal Memory vs. NavigationNoneNone
Verbal Memory vs. Video +NavigationNoneNone
Navigation vs. Video +Verbal MemoryNoneNone
Temporal encodingVideoNoneNone
Verbal MemoryNoneT1>T2: Left LOC (Z=3.97, MNI: −40,–62, 32)
NavigationNoneNone
Verbal Memory vs. VideoNoneNone
Navigation vs. VideoNoneNone
Verbal Memory vs. NavigationNoneNone
Verbal Memory vs. Video +NavigationNoneNone
Navigation vs. Video +Verbal MemoryNoneNone
RetrievalVideoNoneNone
Verbal MemoryNoneNone
NavigationNoneNone
Verbal Memory vs. VideoNoneNone
Navigation vs. VideoNoneNone
Verbal Memory vs. NavigationNoneNone
Verbal Memory vs. Video +NavigationNoneNone
Navigation vs. Video +Verbal MemoryNoneNone
Spatial RetrievalVideoNoneNone
Verbal MemoryNoneNone
NavigationNoneNone
Verbal Memory vs. VideoNoneNone
Navigation vs. VideoNoneNone
Verbal Memory vs. NavigationNoneNone
Verbal Memory vs. Video +NavigationNoneNone
Navigation vs. Video +Verbal MemoryNoneNone
Temporal RetrievalVideoNoneNone
Verbal MemoryNoneNone
NavigationNoneNone
Verbal Memory vs. VideoNoneNone
Navigation vs. VideoNoneNone
Verbal Memory vs. NavigationNoneNone
Verbal Memory vs. Video +NavigationNoneNone
Navigation vs. Video +Verbal MemoryNoneNone

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  1. Li Zheng
  2. Zachary Boogaart
  3. Andrew McAvan
  4. Joshua Garren
  5. Stephanie G Doner
  6. Bradley J Wilkes
  7. Will Groves
  8. Ece Yuksel
  9. Lucia Cherep
  10. Arne Ekstrom
  11. Steven M Weisberg
(2025)
Newly trained navigation and verbal memory skills in humans elicit changes in task-related networks but not brain structure
eLife 14:RP106873.
https://doi.org/10.7554/eLife.106873.3