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Reviewer #1: The authors did a good job of incorporating prior research, theory, and methodology into their approach. The manuscript was well written, and the figures worked well to display the findings. The authors examined whether pattern completion and/or pattern separation are impaired in older adults versus younger adults using several complimentary tasks: the Mnemonic Similarity Task (visual), and two novel tasks: Verbal Discrimination Task (verbal) and the Verbal Completion Task (verbal). As expected, based on prior similar studies using the MST, they found that older adults exhibited impaired “Old”-Lure proportions, Target vs Lure d’, and Lure vs Foil d’, but intact “Old”-Target proportions, “Old”-Foil proportions, and Target vs Foil d’. Importantly, there was a significant interaction between age group and d’ type (Target-Lure d’ vs. Lure-Foil d’). This indicated that the older adults were more likely to treat lures like targets (and less like foils) than the younger adults. For the VDT, they found impaired “Old”-Lure proportions, “Old”-Foil proportions, Target vs Foil d’, and Target vs Lure d’, but intact “Old”-Target proportion and Lure vs Foil d’. Importantly, there was no impairment in Lure vs Foil d’, suggesting that memory specificity was intact in the semantic domain. However, neither younger nor older adults were able to discriminate lures from foils in the VDT (floor effect), making it difficult to determine whether there was an age-related impairment in memory specificity. Direct comparisons across groups between the Lure vs Foil d’ for the MST and VDT revealed more age-related differences for the MST than the VDT. Follow-up analyses of the effect of language familiarity or semantic-similarity on performance did not appear to be relevant the group comparisons of memory specificity in the semantic domain. For VCT, they found impaired Inclusion proportion as well as impaired estimates of automatic and controlled retrieval, whereas the Exclusion proportion was intact. There was no significant correlation identified between the VDT slope (thought to reflect pattern separation) and the automatic estimate from the VCT (thought to reflect pattern completion).

Concerns:

1. The authors found that older adults exhibited different levels of memory specificity on the MST than the VDT (i.e., lure vs foil d’) and stated “This suggests that the effect of age on performance depends on the task type, where greater declines emerge on the Object MST”. This conclusion is difficult to support given the floor effects for the Foil vs Lure d’ for both groups on the VDT. If both groups were above chance on the VDT and the same significant interaction emerged, this would be strong evidence for their claim. It appears they have created a task in which it is impossible for humans to discriminate between lures and foils, which is the critical type of discrimination needed to justify the claim they are trying to make about memory specificity being differentially impaired across tasks in older adults, i.e., that memory specificity is not a domain-general type of impairment that occurs during aging.

2. For the results supporting the analyses in Figure 5, the authors note there was no significant relationship between independent estimates of pattern completion from the VCT and pattern separation from the VDT. Yet, it is possible that these two estimates are uncorrelated because they are from two different tasks. It would be stronger evidence for their claim if they could show that independent estimates of pattern separation from both tasks were significantly correlated and/or that independent estimates of pattern completion from both tasks were significantly correlated. Against that backdrop, their finding that pattern separation and pattern completion were uncorrelated would be more convincing.

Minor Concerns:

1. Abstract: Words like “change” and “decline” are used, but given the non-longitudinal, group-based comparisons these types of words are not accurate. The results should be described in terms of deficits or impairment.

2. Page 17, line 343: “confirming desirable features of the MST design in our sample”. What desirable features are the authors referring to?

3. Page 19, line 389: “This conclusion would be premature however, because our manipulation of semantic similarity was not as pronounced as the object similarity manipulation in the MST”. Please make it clearer how the authors are making the determination that semantic similarity was not as pronounced as on the object similarity in the MST. It is not clear what characteristics or findings they are looking at to make that decision.

4. Page 29, line 610. The authors state that a limitation of their behavioral study was that it did not include neuroimaging data. Reporting on behavior alone within a conceptual framework, but without accompanying neuroimaging data, is not a limitation.

Reviewer #2: ## Summary

The present article, "Aging and episodic memory specificity: Evidence challenging a domain-general pattern separation decline" explores (1) the loss of specificity in healthy aging, found in visual episodic memory, for conceptual similarity and (2) how pattern separation and pattern completion are interdependent for this situation. The study is based on three experimental tasks, a classic mnemonic similarity task, an adapted version manipulating semantic similarity and a verbal completion task to assess semantic pattern completion.

The topic is timely and relevant. Increasing data support the loss of specificity for visual episodic memory. Yet, little is known for conceptual material. In the same vein, it is supposed that pattern completion is interdependent with pattern separation, but behavioral results remain mixed.

Overall, the article is well written, based on an original and adapted methodology, and proposes an interesting contribution to the topic. The introduction offers a comprehensive review of the literature. The experimental paradigms are well designed. The statistical analyses are correctly done and the conclusion is consistent with the data.

Nonetheless, I will have relatively major comments (see below) that should be addressed leading to a major revision.

## Major Comments

One of the study’s aims, stated in the introduction section, is “3) We accounted for other cognitive processes, such as executive functioning or visual-spatial processing”, but no mention of the objective is made in the abstract. I suggest adding this information there.

I am also confused about the control mentioned. The authors used a PDP (process dissociation procedure) procedure, which is suitable to account for the attentional and controlled process, but it is unclear how the visual-spatial processing is taken into account. No further mention of visuo-spatial processing is done in the article. This mention should be removed or relevant data should be added.

Moreover, it is stated that “we administered a neuropsychological test battery to older adults only”, but that these data will not be discussed here. Nonetheless, executive and visuo-spatial results from the neuropsychological assessment should be used to account for the third objective of the paper. By the way, please add, at least as supplementary data, the list of the tests used.

On a statistical level, the age range is very large for the older adults considering that aging decline accelerated around the 70’s. The analysis method should control for it. In the same idea, the education level of the older adults is very high and significantly different from the one in the young adults group. This variable should also be controlled.

I would recommend exploring the correlation matrix between all variables and/or to use, at least ANCOVA, or better, hierarchical regressions or generalized linear mixed as used after (by the way, a yes/no recognition task is more like a Bernoulli distribution than a normal distribution which shouldn’t be processed by ANOVA, see Dixon, 2008).

Some statistical details are also missing such as the use of bilateral or unilateral p-values tests, the control of outliers for accuracy and reaction times, other assumptions checks (normality), and so on.

Please comment on the possible bias related to intentional memorizing content in the semantic adaptation of the MST vs. the incidental learning done in the standard MST.

Perceptual information is richer than conceptual ones. As analogous signals, perceptual information are filled with details (e.g. color nuances, resolution, context, points of view, and so on) that are not presented in a verbal concept (there is a multiple of possible representations of an ELEMENT). Consistent with this idea, phonological similarity induces a similar loss of specificity. Consequently, it is difficult to conclude that the interaction between age and the nature of the stimuli is supported by a preserved processing of the conceptual information. One could imagine that the effect is mainly driven by the degree of details convey in the stimuli used (see also the sentence l. 390 “semantic similarity was not as pronounced as the object similarity manipulation in the MST”). Could the authors better justify their conclusions or acknowledge this hypothesis as a limit of their study. Otherwise, please add more nuanced to the discussion (e.g. “our results add to emerging evidence that age-related decline in memory specificity is restricted to perceptual domains...” ).

## Minor Comments

- All the data should be uploaded on the OSF as dropbox is a suitable and durable sharing method (the data remain on the folders of someone specifically and the access could be easily removed).

- The pictures should be uploaded with higher quality (e.g. 600 DPI).

- Update the abstract to mention the aim to add the classical MST task for a subset of participants (i.e. direct comparison of visual vs semantic similarity, see l).

- For the same reason, update the introduction section presenting the aims of the study (l-139) to explicitly present the rational to “We additionally included the standard object MST for comparison with the VDT for a subset of participants”. If it is an objective of the study, it should be presented with the other ones (l. 125-131) and if it a secondary or exploratory aims, it should be mentioned as it (see l. 210 “Midway through data collection, we became interested in comparing performance in the VDT with the visual object MST.”).

- Please add more details about the power analysis. Was it computed for a simple t-test/ANOVA test or did it account for the hierarchical levels or possible covariables.

- Please provide more details on the ethical approval (e.g. approval id or number).

- In general, please add Notes to the tables to provide details on the acronyms and abbreviations (SD, F, M, MST...). Please also add where the group comparisons were significant.

- I would not agree with the following statement (l. 206-208) “this way, it is more likely that any observed age-related memory impairments are driven by mnemonic changes rather than visual acuity or perceptual changes.”. The pattern separation literature is heavily based on representational-hierarchical approach which did not consider a difference in nature between memory and perceptual processes (see Kent et al., 2016; see also embodied view of cognition Mille et al., 2021).

- It is odd to mix up separated at a similar level the verbal stimuli and the tasks using them. I suggest using a hierarchical structure with something like “Semantic tasks” at a higher level and within it “Verbal stimuli”, “Intentional Encoding Task”.

- L. 321-325 : the paragraph is mainly a redundant part of the method, please remove it.

- Please provide the exact formula used to compute the discriminability index (l. 329).

- Letters used in mathematical expressions should be in italic (F, t, p...) like in F(1,132) = 333.34, p < .

- Please change the verb “tended” (l. 386) as it could imply that the effect is not really significant.

- Please remove this statement for the results “semantic similarity was not as pronounced as the object similarity manipulation in the MST.” or provide more explanations about it, here.

- Please use consistent terminology between semantic and domain-general terms (e.g. l. 522).

- What the authors mean by "reflect a lack of interest in perceptual details” (l. 539-540)?

- I would recommend being more nuanced about “a robust deficit in automatically recall memories” (l. 584) as numerous studies have reported the opposite and as the tasks constraints should be taken into account (e.g. trigram completion has been reported as being more demanding than a real automatic memory recall and to involve some executive components).

## References

- Dixon, P. (2008). Models of accuracy in repeated-measures designs. *Journal of Memory and Language, 59*, 447–456.

- Kent, B. A., Hvoslef-Eide, M., Saksida, L. M., & Bussey, T. J. (2016). The representational–hierarchical view of pattern separation : Not just hippocampus, not just space, not just memory? _Neurobiology of Learning and Memory_, _129_, 99‑106.

- Mille, J., Brambati, S. M., Izaute, M., & Vallet, G. T. (2021). Low-Resolution Neurocognitive Aging and Cognition : An Embodied Perspective. _Frontiers in Systems Neuroscience_, _15_, 687393.

Reviewer #3: This study investigates the impact of semantic relatedness on age differences in pattern separation and completion. The authors designed two new tasks, namely the verbal discrimination and completions tasks. A subset of participants also completed the standard version of the mnemonic similarity task (MST). The results showed that the age-related decline in memory specificity is stronger for perceptual than for semantic relatedness. Older adults were also impaired in pattern completion. The authors interpret these results as evidence against the view that typical aging is associated with a domain-general decline in pattern-separation abilities and a bias toward pattern-completion.

The study question is relevant and topical. Also, the effort the authors have made to design the new tasks, and to characterize the semantic relatedness of all list words, are remarkable. The results of this study will be informative for the cognitive aging scientific community. However, there are some limitations that need to be addressed, which I list below.

1. The design of the present study includes some specific features that must be considered. First, the encoding procedure used for the VCT and VDT is likely to have oriented participants toward a deeper conceptual processing of the target words. Given that numerous previous studies have established that older adults benefit from semantic (and schematic) support, such an encoding strategy may have been especially beneficial to this group compared with younger adults, tempering age differences. Unless the authors disagree with this view, it should be acknowledged in the manuscript. Second, encoding was explicit in the VCT and VDT but incident in the MST; could the authors explain this methodological choice, and how this is considered in the interpretation of the results? Finally, from my understanding, there was no delay between encoding and test in the VCT whereas there was a 10-minute delay in the VDT – is that correct? If so, could the authors also explain this choice and its potential influence on the results?

2. I believe that the authors could further discuss the implications of their results for the nature of memory representations in typical aging. For example, the concept of “gist” is insufficiently developed. The authors mention that older adults tend to rely more on gist memory (p. 3, lines 55-56), but do not define gists. Some relevant references could be cited in the introduction and discussion, such as Grilli & Sheldon (2022), and/or Greene & Naveh-Benjamin (2023), and/or Brainerd & Reyna (2015). Another hypothesis is that memory representations in healthy older adults are “cluttered”, that is, they include more irrelevant and no-longer relevant features, in addition to relevant features, which harms performance in memory tasks that require memory representations of high fidelity (Amer et al., 2023).

Grilli, M. D., & Sheldon, S. (2022). Autobiographical event memory and aging : Older adults get the gist. Trends in Cognitive Sciences, 26(12), 1079 1089. https://doi.org/10.1016/j.tics.2022.09.007

Greene, N. R., & Naveh-Benjamin, M. (2023). Adult age-related changes in the specificity of episodic memory representations : A review and theoretical framework. Psychology and Aging, 38(2), 67 86. https://doi.org/10.1037/pag0000724

Brainerd, C. J., & Reyna, V. F. (2015). Fuzzy-trace theory and lifespan cognitive development. Developmental Review, 38, 89 121. https://doi.org/10.1016/j.dr.2015.07.006

Amer, T., Wynn, J. S., & Hasher, L. (2022). Cluttered memory representations shape cognition in old age. Trends in Cognitive Sciences, 26(3), 255 267. https://doi.org/10.1016/j.tics.2021.12.002

3.A. The observation of a trend for age x similarity interaction effect on correct rejections in the VDT in the participants who reported English as first language is important. Figure 3 distinctly shows that in young adults there was no relation between semantic similarity and correct rejections, whereas these two variables are negatively associated in older adults. Considering that this analysis was conducted in a subsample, it is likely to be underpowered. Thus, if this effect existed, it would go against the authors’ conclusions, for example “our results add to emerging evidence that age-related decline in memory specificity is restricted to perceptual domains, compared to semantic ones” (p. 25, lines 535-537), or “In contrast to older adults’ mainly preserved memory specificity in the semantic domain” (p. 27, line 580). Unless the authors can argue against this, I believe that these interpretations should be nuanced.

3.B. Related to this, the authors highlight the question of why older adults “do not show superior semantic discrimination” although they have “heightened semantic knowledge” (lines 561-562, p.26). It is important to keep in mind that the tasks designed in this study and those used in most similar studies (including that of Ilyés, Paulik, and Keresztes, 2024, SciRep), rely on episodic memory, although the conceptual similarity of targets and lures can be manipulated. Therefore, performance on these tasks is impacted by the fidelity of episodic memory representations, which declines with typical aging. This could partially explain why older adults’ performance on such tests is impaired although they have enhanced semantic knowledge. What do the authors think about this tentative interpretation?

4. Can the authors specify explicitly on what data the multiple linear regression model that aimed at comparing age differences between the object MST and VDT was conducted (p.21, lines 433-442)? My question aims to confirm that the dependent variable here corresponds to memory discrimination, since the authors report an age x task type interaction. If the dependent variable did not correspond to a mnemonic discrimination measure, then a three-way interaction (i.e., age x task type x stimulus, e.g., lure/foil) would be warranted to ensure that the effect can be interpreted as reflecting memory specificity rather than overall memory performance. Line 436 seems to indicate that, but making it explicit could be helpful for future readers.

5. As highlighted by the authors p.27, the use of verbal materials and synonyms to manipulate semantic relations between stimuli is relevant but also comes with limitations. Mentioning ways to overcome such limitations in future studies could be helpful to readers. For example, some researchers have manipulated conceptual similarity for visual material (e.g., Frick et al, 2023; Naspi et al, 2021). Alternatively, the material used by Ilyés et al., 2024 also seems to overcome this limitation, as the related/unrelated lure manipulation comes from the rearrangement of word pairs (e.g., “police uniform” and “official uniform”).

Frick, A., Besson, G., Salmon, E., & Delhaye, E. (2023). Perirhinal cortex is associated with fine-grained discrimination of conceptually confusable objects in Alzheimer’s disease. Neurobiology of Aging, 130, 1‑11. https://doi.org/10.1016/j.neurobiolaging.2023.06.003

Naspi, L., Hoffman, P., Devereux, B., Thejll-Madsen, T., Doumas, L. A. A., & Morcom, A. (2021). Multiple dimensions of semantic and perceptual similarity contribute to mnemonic discrimination for pictures. Journal of Experimental Psychology. Learning, Memory, and Cognition, 47(12), 1903‑1923. https://doi.org/10.1037/xlm0001032

Minor comments

• I am not sure that the study by Delarazan et al. (2023, Learning and memory) cited p.4 (lines 92-94) can be described as investigating “perceptual and semantic” discrimination. Rather, the authors tested memory for narrative details, that is, “moments described” (i.e., how the story unfolded). While I acknowledge that this is close to the semantic domain, it must not be confused with manipulations of conceptual similarity such as the one in the present study or that of Ilyés et al., 2024.

• P.10, line 204: “PC and PS” these abbreviations were not defined in the article.

• P.9, lines 130-131 or lines 136-137, I suggest citing also the study of Gellersen et al. (2021) to support the authors’ argument.

Gellersen, H. M., Trelle, A. N., Henson, R. N., & Simons, J. S. (2021). Executive function and high ambiguity perceptual discrimination contribute to individual differences in mnemonic discrimination in older adults. Cognition, 209, 104556. https://doi.org/10.1016/j.cognition.2020.104556

• p.8, lines 181-182, is the 95% CI of 4.77-6.96 the CI around the t-test? If so, it is surprising that this interval does not include the t value itself (i.e., 10.59). Can the authors double check and correct if needed?

• p.23, lines 481-482 and lines 489-490 seem to repeat the same information (i.e., that VCT performance was not influenced by English language background).

• P.27, line 581, it is unclear what “in their ability to recall memories” refers to, especially since there was no proper memory recall task in this study. Also, the sentence lines 583-584 is almost identical to the first sentence of this paragraph (lines 580-581).

• p.11, line 248, there seems to be a typo in “Based off of these cues”.

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Reviewer #3: No

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Dear Dr. Youm,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

Please submit your revised manuscript by Oct 31 2025 11:59PM. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at plosone@plos.org . When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

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Jie Wang, Ph.D.

Academic Editor

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Reviewer #2: All comments have been addressed

Reviewer #3: All comments have been addressed

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2. Is the manuscript technically sound, and do the data support the conclusions??>

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Reviewer #2: Yes

Reviewer #3: Yes

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Reviewer #2: Yes

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Reviewer #1: The authors have addressed my primary concern about d’ floor effects in the VDT task by explaining that the analysis of VDT slope gets around the floor effects. In addition, the lack of a group effect in the slope analysis is consistent with the lack of a group effect in the d’ analysis. I missed this logic in the initial submission, and it is still missing from the revision. Thus, as a naïve reader of the revision, I would still have the same concern about floor effects that I had upon reading the previous version.

As it stands, the semantic similarity analysis of slopes section begins “The analyses above suggest that the foils in the VDT may have had unintended semantic relationships to studied words. To address this,...”

This introduction to the semantic similarity analysis section is still missing the important issue that has arisen in the prior paragraph, where the floor effect d’ analyses are reported. The authors should make it clearer at this stage of reporting that the slope analysis will address the floor effect issue they have just read about. Also make it clearer in the Discussion that the findings from the murky d’ analysis that suffers from floor effects is nonetheless concordant with the findings from the slope analysis that does not suffer from floor effects.

Finally, given the importance of the slopes for this paper, please report the slope descriptive statistics and p values in Table 2.

All other concerns have been addressed.

Reviewer #2: The authors have done an excellent job to take into account all the comments. The responses and modifications done lead to a valuable article.

Reviewer #3: I commend the authors for the thorough revision of their manuscript. All my comments have been addressed satisfactorily.

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Reviewer #1: No

Reviewer #2: Yes:  Guillaume T Vallet

Reviewer #3: No

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Aging and episodic memory specificity: Evidence challenging a domain-general pattern separation decline

PONE-D-25-21591R2

Dear Dr. Youm,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

Within one week, you’ll receive an e-mail detailing the required amendments. When these have been addressed, you’ll receive a formal acceptance letter and your manuscript will be scheduled for publication.

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Kind regards,

Jie Wang, Ph.D.

Academic Editor

PLOS ONE

Additional Editor Comments (optional):

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

Reviewer #1: All comments have been addressed

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2. Is the manuscript technically sound, and do the data support the conclusions??>

Reviewer #1: Yes

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3. Has the statistical analysis been performed appropriately and rigorously? -->?>

Reviewer #1: Yes

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4. Have the authors made all data underlying the findings in their manuscript fully available??>

The PLOS Data policy

Reviewer #1: Yes

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5. Is the manuscript presented in an intelligible fashion and written in standard English??>

Reviewer #1: Yes

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Reviewer #1: (No Response)

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Reviewer #1: No

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