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The mysterious public career execution of Robert Ryan

On August 24th I received an anonymous email over my website from a „Concerned Microbiologist”:

“I would like to bring the following to your attention on Robert P Ryan at the University of Dundee. https://pubpeer.com/search?q=Robert+P.+Ryan
He holds several high profile research grants and has won several awards. He is under investigation at the University of Dundee.
He has also featured in articles signing his praises in terms of his research achievements.
http://www.scotsman.com/news/university-of-dundee-s-rising-science-stars-1-3932042

Only days after, on August 28th, big news broke out in Scottish and Irish media. The Scotsman brought the headline: “Leading scientist suspended amid ‘research misconduct’ investigation”.  The newspaper then indicated that Ryan has to answer for suspected image manipulations in his papers:

“It is alleged he used identical images across multiple papers, claiming they were different strains. In some cases, it is alleged the evidence was flipped or rotated, which could indicate an “intent to deceive”, according to one source. The extent of the alleged misconduct is unclear, but the source indicated it is alleged to have spanned “a number of years” and involved numerous prestigious journals”.

Similar reports appeared on the same day in The Irish Times, The Courier and Herald Scotland, the latter also wrote:

“It is understood his research group at the university has been dissolved, with PhD students and staff scientists reallocated elsewhere while a formal investigation takes place”.

Robert Ryan is an Irish microbiologist, specializing in bacterial signalling and pathogenesis, who came to work as reader at the Life Science Division of the University of Dundee in Scotland, after a highly successful postdoctoral stay at the lab of leading plant pathogen researcher Maxwell Dow at the University College Cork (UCC) in Ireland. Still with his mentor Dow in Cork, Ryan became principal investigator with his own lab already in 2010 , supported by the Science Foundation Ireland Starting Investigator award and Wellcome Trust. In 2013, Ryan received a faculty position in Dundee, his salary and research are supported for five years by the Wellcome Trust Senior Fellowship. He then won many other awards and research grants, as mentioned in the newspaper articles. In October 2015 Ryan was also appointed Young Investigator by the prestigious life science society EMBO. In brief, Ryan was an absolute success story and a rising star of microbiology and plant pathogen research. Was he actually another dishonest cheater like Olivier Voinnet, luckily stopped short before he became too entrenched in networks and tenure?

okah2ii
Alleged image duplications in Ryan’s papers, source PubPeer user

The PubPeer evidence alleging data duplications in journals like Molecular Microbiology, Journal of Bacteriology, PNAS and EMBO J were posted in March and April 2016. According to newspaper information, Ryan lost his lab before any investigation into his alleged misconduct was performed. Herald Scotland quotes “a university spokesman” from Dundee:

“There is an ongoing investigation into an allegation of research misconduct. In order not to prejudice in any way the outcome of that investigation, we are unable to comment further on it at this time”.

According to Irish Times, this is not a proper in-depth independent investigation, but a brief internal one (which may make its outcome subject to legal appeal):

“Dundee University has launched a formal investigation into the allegations against Dr Ryan, led by university vice-principals. The inquiry is expected to last between two and four weeks”.

Wellcome Trust apparently chose not to scrutinise Ryan’s work, but to rely on the results of the ongoing Dundee investigation, according to Herald Scotland:

“In a statement, the Wellcome Trust added: “Wellcome takes allegations of research misconduct seriously. We expect institutions to investigate any allegation of misconduct, as the University of Dundee are doing in this case, and we would consider taking action only if allegations are upheld”.

What about Ryan old employer University College Cork in Ireland, where all of his PubPeer-flagged papers hail from? Irish Times quotes “a spokeswoman for UCC”:

“UCC understands that there has been an allegation of research misconduct in relation to a member of staff at the University of Dundee. […] Should any action be warranted by UCC in due course, UCC will follow up at that point.”

To sum up, the available information indicates Ryan lost his lab and had his career destroyed on suspicion alone. He will not even receive a proper independent investigation, it seems. At the same time, it is highly unlikely that four regional newspapers would coincidentally follow up and bring this story simultaneously, based solely on some PubPeer evidence. The fact that Ryan was delivered for public crucifixion by the mass media indicates that a senior source from inside University of Dundee has briefed the newspapers officially. I wonder therefore who contacted me over my website.

Obviously, everyone involved received a gag order. Ryan did not communicate with any of the newspapers. Dow, Ryan’s mentor of many years and key co-author on all Ryan’s PubPeer-flagged publications, only replied to my inquiry with:

“I am afraid I cannot comment on this issue at this time”.

robertandmax_2
Ryan and his mentor Dow, source UCC

A major scandal, a breach of basic institutional privacy and employee rights guidelines as well as the right to fair trial? Should Ryan get himself a lawyer specialising on labour law and sue University of Dundee for all they have? Given the apparent absence of any investigative verdict, and the immense reputation damage he sustained, his chances to win in court would be practically 100%.

Maybe not. A huge silent elephant in the room has been overlooked, namely the only other major institution with investigative power and a big stake in the Ryan scandal: EMBO.

This respected European society recently started funding Ryan as Young Investigator, while one of his questioned papers appeared in 2013 in their own flagship journal, EMBO J. EMBO director Maria Leptin wrote to me on August 25th, 3 days before the news broke in the newspapers:

“We are aware of the allegations, and as usual, are carefully assessing whether actions might need to be taken”.

I contacted Leptin with my inquiry if there was a successful EMBO investigation into Ryan’s case already, but received no reply yet. If Ryan was indeed found guilty of data manipulation elsewhere (with EMBO the strongest candidate), University of Dundee would have an easy case suspending him and even sacking him after a brief internal investigation. Everything would fall into place.

In fact, this was exactly what EMBO did only recently. After they were informed in December 2015 through my article about PubPeer allegations against their recent awardee Sonia Melo, EMBO initiated own investigation. Already by the end of February 2016, the EMBO inquiry was done, with the result that Melo lost her Young Investigator funding and was suspended as PI by her Portuguese employer Instituto de Investigação e Inovação em Saúde (I3S) at the University of Porto. The investigation there is still ongoing, as I was informed by the I3S communication unit coordinator Júlio Borlido Santos on July 29th :

“As far as I am aware, the analysis of Dr. Melo’s situation is still under way, conducted by the designated external committee. As soon as the Communication Unit is informed of the next steps to be taken, public statements will be publicly released”.

Thus, the Ryan case may be actually very similar to that of Melo. The only problem is that EMBO investigative reports are not published and only available to institutions directly involved, such as Melo’s I3S or possibly, Ryan’s University of Dundee. This secrecy is supposed to serve privacy and personal protection of the accused. Instead, if Ryan was really investigated by EMBO, this false confidentiality achieved exactly the opposite.

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Alleged image duplications in Ryan’s paper. Source PubPeer user.

Update 14.11.2016. the shroud of mystery is slightly lifting, please read here.

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23 comments on “The mysterious public career execution of Robert Ryan

  1. Anonymous

    I used to work in the same dept. I had no idea. I am also very surprised that this leaked as it has. No previous issues with staff have come out like this (issues are inevitable in any research department, this is not to imply anything in particular) and even the most senior egos are on brand and on message. It is also, generally, a supportive workplace.

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  2. If he has been suspended and group disbanded, as well as having more than ten entries in his honor on pub peer it will be fairly common knowledge in his building/research area. Seems a big leap to think that the University briefed the press instead of someone just stirring the pot.

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  3. Ana Pedro

    In cases like this one and Sonia Melo’ s case both first and seniors authors as well as other contributing authors should be fully investigated in order to be fairly judged.

    Like

  4. It seems to me that the University of Dundee will not easily decide to suspend a famous researcher like Robert Ryan and it seems to me that such a suspension is an interesting news item for readers of a Scottish newspaper.

    It also seems to me that a Scottish / Irish / UK journalist who becomes aware of this suspension will conduct his own investigation to sort out the details / backgrounds why this famous researcher got suspended. No one has until now indicated stated that the suspension of Robert Ryan is not true and no one has until now indicated / stated that this suspension relates to something totally different (issues which have nothing to do with the allegations that Robert Ryan “used identical images across multiple papers”).

    I fail to understand why Robert Ryan and/or one or more of the (many) co-authors of for example the paper in EMBO J (see https://pubpeer.com/publications/23881098 ) have until now not rebutted the comments of peer 1 (posted on 19 April 2016). This paper counts in total 11 authors, so why not rebut the views of peer 1 about the similarities in the depicted leaves?

    The same is of course the case for for example the comment of peer 1 (also posted on 19 April 2016) about leaves in a 2006 paper in Molecular Microbiology (see https://pubpeer.com/publications/B1BAB266DC2527F429653D32BC8BB6 ). This paper counts in total 8 authors.

    So I can imagine that the University of Dundee has told to Robert Ryan that he is at the moment not allowed to contact journalists / others / other scientists, but quite a few (many?) of his co-authors will not have an affilation to the University of Dundee. These researchers are thus free to rebut the claims of for example peer 1 about the depicted leaves in these papers (see above).

    So why has until now no one of all these authors not rebutted the views / statements of peer 1 about the depicted leaves in these two papers?

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  5. Jaime A. Teixeira da Silva

    There is no mystery here.

    I made a formal inquiry (see email below) to several British research institutes on April 20, 2016. This followed the registration of possible concerns or errors at PubPeer, and also documentation at Retraction Watch on April 19, 2016:
    http://retractionwatch.com/2014/01/25/weekend-reads-trying-unsuccessfully-to-correct-the-scientific-record-drug-company-funding-and-research/#comment-1011411
    http://retractionwatch.com/2014/01/25/weekend-reads-trying-unsuccessfully-to-correct-the-scientific-record-drug-company-funding-and-research/#comment-1011416
    http://retractionwatch.com/2014/01/25/weekend-reads-trying-unsuccessfully-to-correct-the-scientific-record-drug-company-funding-and-research/#comment-1011424
    http://retractionwatch.com/2014/01/25/weekend-reads-trying-unsuccessfully-to-correct-the-scientific-record-drug-company-funding-and-research/#comment-1011434

    This is the email I addressed to Professor J. Maxwell Dow, Dr. Ryan and other co-authors (emails redacted, but are available upon request):

    “J. Maxwell Dow: 6X Xanthomonas campestris queries

    Prof. J. Maxwell Dow
    BIOMERIT Research Centre, Department of Microbiology, BioSciences Institute, University College Cork, Cork, Ireland
    http://publish.ucc.ie/researchprofiles/D010/mdow
    Dr. Robert P. Ryan
    Division of Molecular Microbiology, College of Life Sciences, University of Dundee, Dundee, United Kingdom
    http://www.lifesci.dundee.ac.uk/people/robert-ryan
    Prof. Jane Rogers (+ Dr. Leah Clissold)
    Former Director, The Genome Analysis Centre, Norwich Research Park, Colney Lane, Norwich, United Kingdom
    http://cnrgv.toulouse.inra.fr/Communication/Plant-Genomes-Day-2014/Scientific-Program/Jane-Rogers
    Dr. Melanie Febrer
    Next Generation Sequencing Core Facility, University of Dundee, Dundee, United Kingdom
    http://www.lifesci.dundee.ac.uk/groups/irwin_mclean//resources
    Dr. Maxwell (de Oliveira) Andrade
    Citrus Research and Educational Center, Department of Microbiology and Cell Science, University of Florida, USA
    https://www.researchgate.net/profile/Maxuel_Andrade

    Dear Prof. Maxwell Dow et al.,

    I follow issues in plant science publishing quite closely and read the plant science literature very widely. Your research and papers are important for the phytopathology community, and are of interest to me.

    Could you be so kind as to offer some commentary about your 6 papers, please, and indicate a time-line by which you plan to respond and correct the literature, where required. The editors and publishers have not yet been contacted, with the hope, in good faith, that you and your colleagues will address all of the issues raised publicly.

    68.1

    An, S.-Q., Febrer, M., McCarthy, Y., Tang, D.-J., Clissold, L., Kaithakottil, G., Swarbreck, D., Tang, J.-L., Rogers, J., Dow, J. M. and Ryan, R. P. (2013a), High-resolution transcriptional analysis of the regulatory influence of cell-to-cell signalling reveals novel genes that contribute to Xanthomonas phytopathogenesis. Molecular Microbiology, 88: 1058–1069.
    doi: 10.1111/mmi.12229
    http://onlinelibrary.wiley.com/doi/10.1111/mmi.12229/epdf
    http://onlinelibrary.wiley.com/doi/10.1111/mmi.12229/abstract
    https://pubpeer.com/publications/50B2EB760523D9573A3E161E021091

    68.2

    Shi-Qi An, Ko-Hsin Chin, Melanie Febrer, Yvonne McCarthy, Jauo-Guey Yang, Chung-Liang Liu, David Swarbreck, Jane Rogers, J Maxwell Dow, Shan-Ho Chou, Robert P Ryan (2013b) A cyclic GMP-dependent signalling pathway regulates bacterial phytopathogenesis. The EMBO Journal 32(18), 2430-2438.
    DOI: 10.1038/emboj.2013.165
    http://emboj.embopress.org/content/32/18/2430

    Click to access 2430.full.pdf

    https://pubpeer.com/publications/99BC8436A5059044A120953295A871

    68.3

    Ryan, R. P., Fouhy, Y., Lucey, J. F., Jiang, B.-L., He, Y.-Q., Feng, J.-X., Tang, J.-L. and Dow, J. M. (2007), Cyclic di-GMP signalling in the virulence and environmental adaptation of Xanthomonas campestris. Molecular Microbiology, 63: 429–442.
    doi: 10.1111/j.1365-2958.2006.05531.x
    http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2958.2006.05531.x/abstract
    http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2958.2006.05531.x/epdf

    68.4

    MCCARTHY, Y., RYAN, R. P., O’DONOVAN, K., HE, Y.-Q., JIANG, B.-L., FENG, J.-X., TANG, J.-L. and DOW, J. M. (2008), The role of PilZ domain proteins in the virulence of Xanthomonas campestris pv. campestris. Molecular Plant Pathology, 9: 819–824.
    doi: 10.1111/j.1364-3703.2008.00495.x
    http://onlinelibrary.wiley.com/doi/10.1111/j.1364-3703.2008.00495.x/abstract
    http://onlinelibrary.wiley.com/enhanced/doi/10.1111/j.1364-3703.2008.00495.x

    https://pubpeer.com/publications/B1BAB266DC2527F429653D32BC8BB6

    68.5

    PNAS April 25, 2006 103(17): 6712–6717
    Cell–cell signaling in Xanthomonas campestris involves an HD-GYP domain protein that functions in cyclic di-GMP turnover
    Robert P. Ryan, Yvonne Fouhy, Jean F. Lucey, Lisa C. Crossman, Stephen Spiro, Ya-Wen He, Lian-Hui Zhang, Stephan Heeb, Miguel Camara, Paul Williams, J. Maxwell Dow
    DOI: 10.1073/pnas.0600345103
    http://www.pnas.org/content/103/17/6712

    Click to access 6712.full.pdf

    https://pubpeer.com/publications/54D08BB40C04AB4CAEF659CD7A13C8

    68.6

    PNAS March 30, 2010, 107(13): 5989–5994
    Cell–cell signal-dependent dynamic interactions between HD-GYP and GGDEF domain proteins mediate virulence in Xanthomonas campestris
    Robert P. Ryan, Yvonne McCarthy, Maxuel Andrade, Chuck S. Farah, Judith P. Armitage, J. Maxwell Dow
    DOI: 10.1073/pnas.0912839107
    http://www.pnas.org/content/107/13/5989

    Click to access 5989.full.pdf

    https://pubpeer.com/publications/F0DDBA55ED5EAC7DD3036BD91C1F91

    Many thanks in advance. Kindly acknowledge this email.

    Jaime A. Teixeira da Silva
    https://www.researchgate.net/profile/Jaime_Teixeira_Da_Silva
    Disclaimer: Independent scientist not affiliated with any blog or institute.”

    This was the response I received from Prof. Dow:
    “On Wednesday, April 20, 2016 5:42 AM, “Dow, Max” [redacted] wrote:
    Dear Dr Teixeira da Silva,
    Thank you for your message. We can assure you that we intend to address issues arising with these papers and correct the literature where appropriate as expeditiously as possible.
    Best wishes,
    Max Dow (on behalf of all authors)”

    Of direct relevance that suggests that other very powerful co-authors in very powerful positions may have made Robert P. Ryan the sacrificial lamb:
    http://www.tgac.ac.uk/news/35/15/TGAC-Director-to-step-down/

    There is certainly much more to this than meets the eye.

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  6. “At the same time, it is highly unlikely that four regional newspapers would coincidentally follow up and bring this story simultaneously, based solely on some PubPeer evidence”
    The Irish Times is not a regional paper, it is a national paper. That, indeed, is all the more worrying.

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  7. Scientists are not beyond temptation. Just Google scientific fraud is rife. Why is there such a surprise if someone had made up data? It takes a brave University to investigate these matters properly and many Institutions would cover the matter up and even find a scapegoat. The frailty of the scientific peer review process has been recognized on two occasions by the UK Science Select Committee. They wanted the creation of a science oversight body to take overall responsibility and they were defied by RCUK and UUK. A truly independent investigation would happen with other professions such as medical doctors and it would be carried out by peers. This takes decisions away from local HR Departments who are not equipped for these matters. Scientific publication cannot possibly be considered as confidential yet HR people will try and make the matter confidential.

    The sooner we have a professional body within the UK, with responsibility of investigating science malpractice the better. The Royal Society does not fulfill such a role, indeed it stubbornly refuses to recognize the problem. Its obvious from this case alone that a number of individuals and a number of institutions have been involved creating a situation where everybody can blame everybody else. An overarching body would not be limited by corporate politics.

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  8. Stephanie Cool

    Some further information – It would appear that these allegations have been investigated and extend beyond gossip – There is also no further information about him on the Dundee website.

    http://retractionwatch.com/2016/11/11/prominent-researcher-in-scotland-resigns-after-misconduct-finding-upheld/

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  10. Many of the work particularly with cyclic DiGMP associated phenotypes and quorum sensing are not repeatable even in their model organism Xanthomonas campestris by Ryan and Max Dow his mentor. Surprisingly the Axa21 story which Dr. Ronald lab retracted due to strain problem was repeated by Dr Ryan and his mentor Max Dow and their collaborators in animal pathogenic bacteria and they repeated the same story. I think that their work need to be repeated as many of their findings have different reports from other groups with not so high profile as Dr Ryan and his mentor and long time collaborator Max Dow published work.
    DS

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    • Teddy Dunphy

      Thank you for your comments on Dr Ryan and Dow’s work. I have worked with both on and off for several years. I would very much like to know what phenotypes you are referring to in your post. I can assure you that the phenotypes associated with DSF and the Rpf cell-cell signaling system are solid and reproducible. They have been reported by numerous groups and strains confirmed and used by other groups. With regards the phenotypes observed in cyclic-di-GMP knockouts strains in Xanthomonas I again can assure you these been seen by a number of labs and the strains confirmed. In particular, phenotypes associated with HD-GYP and PilZ domain knockouts have been seen in most Xanthomonads with various degrees of potency. Importantly, the phenotypes are generally associated with virulence in plants and not seen in vitro making them difficult to detect.

      Finally, the major reason for the retraction of these papers from the Ronald laboratory were the absence of a specific effect of Ax21 on plant defence induction in rice lines carrying the Xa21 gene. In addition, it was shown that Ax21 secretion involved type II secretion and not the type I system encoded by rax genes, as initially thought. This was reported in a correction to one of these papers (the peerJ paper). It was not questioned whether the extracellular perception of Ax21 (or derived peptides) had a role in alteration of gene expression in Xanthomonas, as distinct from plants. In the experiments on Stenotrophomonas by the Ryan and Dow, there were no experiments on effects of peptides/proteins on plants and the mutation of a homolog of a gene encoding the type I secretion protein had only a slight effect on the Ax21 level. Importantly, these effects were measured indirectly as the effects of supernatants on the ax21 mutant in S. maltophilia. This point was raised in the peerJ paper, although without mentioning that the mutation still had significant signaling activity in the supernatant. Thus the conclusions seem to remain valid. This was discussed in detail at the last XGC meeting two years ago. I do think that if there are errors in the Stenotrophomonas article it will retracted.

      Why not contact the authors and ask for input? Or contact the Journal and ask to write a common?

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      • Thanks for your comments.
        I am enclosing the comments from Pamela Ronald paper and her comments from the retraction watch which clearly describes that Ryan et al. data is a repetition of a already retracted paper which the author (PC Ronald) honestly self retracted and described how Ran and Dow Axa21 story has lot of problem.
        Furthermore, more soon details of the DSF phenotypes and cyclic DiGMP related problems will be discussed (Hint” DSF promotes biofilm dispersal ?).
        https://peerj.com/articles/242/
        Please go through Dr. PC Ronald own article in https://blogs.scientificamerican.com/food-matters/lab-life-the-anatomy-of-a-retraction/)
        Quote from the Article
        “We were ecstatic to identify this bacterial protein because it provided an important piece to the puzzle of how the rice plant is able to respond to infection. News of the discovery rippled through our community of researchers and drew others into the investigation.
        For example, shortly after we reported our findings at a small meeting of Xanthomonas researchers, another laboratory (Axa21 story of Ryan and Dow) rushed to publish a paper reproducing key aspects of our results in a human pathogen. Soon after, three more papers reproducing our results were published.
        Yet, in a bizarre twist, just as these other laboratories were validating our research, we were discovering reasons to question our own findings”

        Sections from the Peer J paper: https://peerj.com/articles/242/
        Subsequent research from our lab, focused on the biological function of Ax21 in Xanthomonas, suggested that Ax21 serves as a cell–cell signaling molecule mediating motility, biofilm formation and virulence (Han et al., 2011). However, this report was also retracted (Ronald, 2013; Han et al., 2013). Here too, several labs have reproduced some of our results in other bacterial species (McCarthy, Dow & Ryan, 2011; Wang et al., 2013; Qian et al., 2013). For example, McCarthy, Dow & Ryan (2011), investigating the role of Ax21 in S. maltophilia, reported that Ax21 secretion depends on RaxB and that Ax21 serves as a cell–cell signaling molecule involved in motility, biofilm formation and virulence. Qian et al. (2013) identified Ax21 in a proteomic study of the Xoc RS105 secretome. They showed that deletion of the ax21 gene resulted in reduced biofilm formation and extracellular-polysaccharide production.
        These results conflict with publications by two independent research groups. The first, by McCarthy, Dow & Ryan (2011), showed that Ax21 activity from supernatants of S. maltophilia strain K279a carrying a mutation in a presumed raxB ortholog (Smlt2001) is reduced compared with that of supernatants from the wild type strain. From this experiment the authors concluded that RaxB is partially required for Ax21 secretion. However, our analysis of Smlt2001 reveals that it lacks the unique N-terminal peptidase domain (C39, residues 17-143 in the PXO99A genome annotation), characteristic of the RaxB ABC-transporter. Instead, Smlt2001 shows a much higher similarity to PXO_01193, a PXO99 ABC-transporter that lacks the N-terminal C39 peptidase domain. This analysis suggests that Smlt2001 is not an ortholog of RaxB. We were not able to identify a predicted RaxB ortholog in the S. maltophilia K279a genome. The second publication, by Wang et al. (2013), reported that Ax21 is not secreted in the absence of RaxA in Xoc. We have not tested the role of RaxA in Xoc.

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  12. Teddy Dunphy

    Dear DS,

    Thanks for the reply. To address your points:

    I agree there is some conflict in the data between groups working in Ax21. However, as I mentioned regarding the Ax21 story there are serious differences between the Ronald work done in Xanthomonas and the work by Dow in Stenotrophomonas. I am very aware of the PeerJ paper as I pointed to it in my last post. It has some great points about these complexities. The section of the paper you posts even explains the limitation of the experiments. It explains that Stenotrophomonas does not have a conventional rax system for secretion, there are two homologs of Ax21 and that all the experiments carried out by Dow used culture supernatants.

    With regards the other problems, making mysterious statements like “DSF promotes biofilm dispersal?” is not very helpful. So it is difficult make any statement there. All I can say is that it has been clearly seen in a lot of manuscripts that DSF under various different conditions alters biofilm in different ways both causing biofilm formation as well as dispersal. This is dependent on the bacterium and the environment.

    It is clear I am not going to convince you. I suppose that is science. I would ask that you again to contact the authors and ask for input? Or contact the Journal and ask to write a comment, which the authors can respond to maybe?

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  13. Elizabeth Wager

    According to newspaper reports, the University of Dundee has notified affected journals, but without a public report of their investigation readers are left in the dark about how many articles are affected and whether the university has examined all of Ryan’s output, or just some of it. As Leonid has noted, most of the details of this case seem to have come from leaks. Surely it would be better (for readers and co-authors) if there was more transparency on this? Does anybody have any idea how many articles are affected (or do we have to try to piece the answers together from PubPeer)?

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