Piper Group - Neural stem cells in development and disease

Research directions

The human brain is an incredibly complex organ, consisting of over 100 billion neurons, and even more glial cells. Further adding to this complexity is the fact that there are a wide variety of distinct neuronal subpopulations within the brain, each with different morphological characteristics, neurochemical properties and patterns of connectivity. Amazingly, nearly all of the cells within the brain are derived from a relatively small population of neural stem cells (NSCs) that proliferate, then differentiate, during embryogenesis. Understanding how NSC biology is coordinated, both spatially and temporally, to generate the mature brain remains one of the great challenges in biology. My vision is to reveal the mechanisms that control NSC differentiation within the developing brain, and to apply this knowledge to understand disease processes caused by abnormal NSC differentiation (Figure 1).

I have made a number of significant contributions to understanding how NSC differentiation is coordinated during neural development since starting my own group in late 2010. This work, which was supported by competitive fellowship (NHMRC Career Development Fellowship 2009-2012; ARC Future Fellowship 2013-2017) and grant funding (three NHMRC project grants as CIA; two ARC Discovery Projects as sole CI), has helped to elucidate the fundamental mechanisms underpinning neurogenesis within the neocortex, hippocampus and cerebellum. I have also defined critical molecular controllers of NSC quiescence, a cellular state that ensures the longevity of adult NSCs, as well as describing the behavioural consequences of aberrant adult neurogenesis. Finally, I have provided new insights into how abnormal stem cell biology can contribute to a range of neurodevelopmental disorders, as well as cancers of the brain and skin. The significance of my findings has been recognised by multiple awards for research excellence, from both national (e.g. 2018 Emerging Leader Award, Australian and New Zealand Society for Cell and Developmental Biology; 2010 AW Campbell Award, Australasian Neuroscience Society) and international agencies (2015 Innovator Award, Hydrocephalus Association; 2010 CJ Herrick Award, American Association for Anatomists). I now am in an ideal position to address aspects of two key questions in the field, namely, what are the transcriptomic and epigenomic factors that control the differentiation of NSCs during brain development, and how do deficits in this process contribute to disease?


Current collaborations

  • Professor Linda Richards – Queensland Brain Institute, The University of Queensland
  • Associate Professor Thomas Burne – Queensland Brain Institute, The University of Queensland
  • Professor Richard Gronostajski – Department of Biochemistry, State University of New York at Buffalo
  • Professor Francois Guillemot – Division of Molecular Neurobiology, National Institute for Medical Research, Mill Hill, London, UK
  • Associate Professor Mikael Boden – School of Chemistry and Molecular Biosciences
  • Dr Aaron Smith – QUT
  • Professor Raj Thakker – Oxford University
  • Dr Fiona Simpson – Diamantina Institute, The University of Queensland
  • Professor Matt Scott – Stanford University


Dr Michael Piper
NHMRC Research Fellow
Queensland Brain Institute and the School of Biomedical Sciences
The University of Queensland
Brisbane Qld 4072
Tel: (+61 7) 3365 4484
Email: m.piper@uq.edu.au

Opportunities for new researchers

Our lab uses a variety of techniques to probe neural stem cell biology, including molecular biology, immunocytochemistry, magnetic resonance imaging and behavioural approaches.  We also use a range of high end imaging modalities.  Our lab comes from diverse backgrounds, and we are always looking for people with complementary techniques, such as electrophysiology or bioinformatics, to join our group.

Positions are available across our research program for undergraduates, honours students, PhD students, and postdocs, and we are happy to entertain new ideas.  If you are interested in joining the lab, please contact Mike.

Understanding the drivers of neural stem cell differentiation

What are the mechanisms that control neural stem cell (NSC) differentiation during embryogenesis, and that enable the generation of the diverse suite of neurons and glia that comprise the brain? This is a key question in developmental neuroscience. My contribution to this field to date has been to reveal central transcriptional regulators that mediate NSC biology within the brain. Using rodent model systems, I demonstrated that transcription factors of the Nuclear Factor One (NFI) family mediate NSC proliferation and differentiation in the embryonic, postnatal and adult nervous system. This work has received international recognition, as evidenced by numerous invited international presentations and high-impact reviews (e.g. Trends in Cell Biology), and forms the framework around which the hypotheses of this program will be addressed.

I am interested in defining how NSC proliferation and differentiation is regulated at a transcriptional and epigenomic level within the developing nervous system. Using the developing mouse brain as a model system, we are using a suite of molecular and cellular techniques to understand how diverse regions of the nervous system are generated, including the cerebral cortex, the cerebellum, the spinal cord and the hypothalamus. For example, within the cerebral cortex, we are investigating how the NFI family of transcription factors mediate NSC differentiation, and how mutations to the NFI family culminate in macrocephaly. Moreover, we are using mice lacking the gene Nsd1 (a histone modifying protein) to investigate the development of a human syndrome known as Sotos syndrome, which is also characterised by macrocephaly. In collaboration with Mikael Boden (SCMB) and Matt Scott (Stanford), we are also investigating how changes to chromatin landscapes mediate NSC differentiation, and developing bioinformatic tools to enhance the analysis of RNA-seq and ChIP-seq datasets. Collectively, this work will provide fundamental insights into neural development, as well as insights into human neurodevelopmental disorders that arise as a result of abnormal neural stem cell biology in utero.

Development of the cerebral cortex
Development of the cerebral cortex
The cerebral cortex is a critical part of the brain.  Here, a coronal section of the developing mouse cerebral cortex is shown.  There are two main progenitor cell types in the developing cortex, the radial glia (shown in red) and intermediate progenitor cells (shown in green). Defects in the proliferation and differentiation of these cells can give rise to cortical deficits such as microcephaly (smaller brain) or macrocephaly (larger brain). Image courtesy of Lachlan Harris.


Adult neurogenesis

The birth of new neurons within the cerebral cortex, a process termed neurogenesis, plays a critical role in learning, memory and spatial navigation. We are investigating various aspects of adult neurogenesis in rodent models, including neural stem cell quiescence and dendrite elongation and spine formation. We are also interrogating the consequences of abnormal neurogenesis using behavioural tests for learning and memory.

We employ a range of transgenic mice to investigate adult neurogenesis, coupled with techniques ranging from immunocytochemistry, behavioural testing, analysis of axonal connectivity and genome-wide sequencing platforms.  Given the critical roles that learning and memory play in our everyday lives, and the fact that neurogenesis within the adult brain diminishes with age, this research will provide fundamental insights into how this vital process is co-ordinated at a cellular and molecular level.

Adult neural stem cells
Adult neural stem cells
Neural stem cells are found within the dentate gyrus of the adult mammalian brain. Here, a neural stem cell in the dentate gyrus of the adult mouse brain has been identified by the expression of GFAP (green), a marker protein expressed by these cells. Cell nuclei are labelled in white (image courtesy of Lachlan Harris).


Immature neurons
Immature neurons
Newly born neurons within the adult dentate gyrus extend processes and become integrated into the existing hippocampal circuitry.  In this image, newly born, immature neurons are identified through the expression of DCX (in green) within the adult mouse dentate gyrus. The new neurons can be seen extending processes into the granule cell layer of the dentate gyrus (cell nuclei can be seen in white). Image courtesy of Lachlan Harris.


Hippocampal neurogenesis
Hippocampal neurogenesis
Neurons born within the adult hippocampus from endogenous neural stem cells play a key role in learning and memory.  Here, a neuron from the adult mouse dentate gyrus (where neurogenesis within the adult hippocampus takes place) is labelled.  The elaborate dendritic tree of the neuron is critical to enable learning and memory to take place. Image courtesy of Sabrina Oishi.


Identifying how abnormal neural stem cell biology contributes to disease

The importance of NSC biology to brain development is underscored by disorders associated with abnormal NSC differentiation, including autism, hydrocephalus and macrocephaly. Despite the role of aberrant NSC development to these disorders, our understanding of the cellular and molecular deficits that contribute to disease onset and progression remains limited. Recently, my work has begun to focus on these disorders. Moreover, as the transcriptional landscape of many cancers resembles that of stem cells during development, I am also applying my expertise to understand how abnormal transcriptional activity contributes to cancer progression. This approach has gained significant traction, as evidenced by international awards (2015 Innovator Award, Hydrocephalus Association) and grants (Simons Foundation Autism Research Initiative, 2018-2019; Cancer Council Queensland, 2016-2017) I have received.

We are using our fundamental understanding of NSC biology to investigate numerous neurodevelopmental disorders. We are investigating macrocephaly (enlarged brain size), with a specific focus on the overgrowth syndromes Malan syndrome and Sotos syndrome. This work is supported by clinical collaborators, including Raj Thakker (University of Oxford). We are also collaborating with Dominic Ng (SBMS, UQ) and Kum Kum Khanna (QIMR) to investigate genetic causes of microcephaly.

Abnormal NSC differentiation can also contribute to autism in human patients. In collaboration with Lachlan Jolly (University of Adelaide) and Peter Penzes (Northwestern University, USA), I have been studying how aberrant protein stability can influence the trajectory of NSC differentiation during embryogenesis. I have a mouse model of abnormal protein deubiquitinylation (Usp9x knockout) that shows abnormal neurogenesis and autistic-like traits, and that mirrors phenotypes seen in human patients with mutations to this gene. We are now using this model, along with iPSC-derived cells, to understand how mutations to USP9X culminate in autism. Finally, NSCs also give rise to ependymal cells at the end of corticogenesis, and abnormal ependymal cell differentiation can lead to hydrocephalus. This is a relatively common disorder but how abnormal ependymal cell development contributes to hydrocephalus has not received significant attention. In collaboration with Helen Cooper (Queensland Brain Institute), we are using transgenic mouse lines and transcriptomic profiling to define the molecules that promote ependymal cell differentiation. Ultimately we envisage that this work will lead to ways in which ependymal cell differentiation from NSCs can be modulated to prevent the onset of hydrocephalus.

Finally, I am using my expertise in development to investigate metastatic spread of cancer. Cancer is often viewed as the reacquisition of a developmental genetic signature by a mature cell. With this in mind, a new direction of my laboratory has been to forge linkages with basic and clinical cancer researchers, in order to apply our knowledge and skills to better understand disease progression. We have been focussing on metastatic migration, as we, and others, have shown that misexpression of NFIB regulates the malignant spread of cancer. We are using two different models to investigate the role of NFIB in metastasis, namely melanoma (in collaboration with Aaron Smith, QUT) and perineural invasion of carcinomas of head and neck (in collaboration with Ben Panizza, Princess Alexandra Hospital, and Fiona Simpson, TRI, with whom I recently shared a pilot grant to establish these techniques)

Cultured melanoma cells
Cultured melanoma cells
Melanoma cells show a large degree of heterogeneity. Here, melanoma cells are grown as spheres in an in vitro culture system. We have used immunocytochemistry to reveal the expression of BRN2 (red) and MITF (green) in these melanoma-spheres, revealing the differential expression of these factors within the cellular population. We hypothesise that BRN2 drives NFIB expression to promote metastasis
 (image courtesy of Mitchell Fane).


Piper M, Gronostajski RM, Messina G (2019) Nuclear Factor One X in development and disease. Trends in Cell Biology 29, 20-30.

Oishi S, Harkins D, Kurniawan N, Kasherman M, Harris L, Zalucki O, Gronostajski R, Burne T, Piper M (2019) Heterozygosity for Nuclear Factor One X in mice models features of Malan syndrome. EBioMedicine, 39, 388-400.

Zalucki O, Harris, L, Harvey TJ, Harkins D, Widagdo J, Oishi S, Matuzelski E, Yong X, Schmidt H, Anggono V, Burne THJ, Gronostajski RM, Piper M (2019) NFIX-mediated inhibition of neuroblast branching regulates migration within the adult mouse ventricular-subventricular zone. Cerebral Cortex, in press.

Harris L, Zalucki O, Clement O, Fraser J, Matuzelski E, Oishi S, Harvey TJ, Burne TH, Heng JI, Gronostajski RM, Piper M (2018) Neurogenic differentiation by hippocampal neural stem and progenitor cells is biased by NFIX expression. Development, 145:1-12.

Matuzelski E, Bunt J, Harkins D, Lim JWC, Gronostajski RM, Richards LJ, Harris L, Piper M (2017) Transcriptional regulation of Nfix by NFIB drives astrocytic maturation within the developing spinal cord. Developmental Biology, 423, 286-297.

Fane ME, Chhabra Y, Hollingsworth DJE, Simmons JL, Spoerri L, Oh TG, Chauhan J, Chin T, Harris L, Harvey TJ, Muscat GEO, Goding CR, Sturm RA, Haass NK, Boyle GM, Piper M, Smith AG (2017) NFIB mediates BRN2 driven melanoma cell migration and invasion through regulation of EZH2 and MITF. EBioMedicine, 16, 63-75.

Harris L, Zalucki O, Gobius I, McDonald H, Osinski J, Harvey TJ, Essebier A, Vidovic D, Gladwyn-Ng I, Burne TH, Heng JI, Richards LJ, Gronostajski RM, Piper M (2016) Transcriptional regulation of intermediate progenitor cell generation during hippocampal development. Development, 143, 4620-4630.

Lim J, Donahoo A, Bunt J, Edwards T, Fenlon L, Liu Y, Zhou J, Moldrich R, Piper M, Gobius I, Bailey T, Wray N, Kessaris N, Poo M, Rubenstein J, Richards L (2015) EMX1 regulates NRP1-mediated wiring of the mouse anterior cingulate cortex. Development,142: 3746-3757.

Heng YHE, Zhou B, Harris L, Harvey T, Smith A, Horne E, Martynoga B, Andersen J, Achimastou A, Cato K, Richards L, Gronostajski R, Yeo G, Guillemot F, Bailey T, Piper M (2015) NFIX regulates proliferation and migration within the murine SVZ neurogenic niche. Cerebral Cortex 25:3758-3778.

Piper M, Barry G, Harvey TJ, McLeay R, Smith AG, Harris L, Mason S, Stringer BW, Day BW, Wray NR, Gronostajski RM, Bailey TL, Boyd AW, Richards LJ (2014) NFIB-mediated repression of the epigenetic factor Ezh2 regulates cortical development. The Journal of Neuroscience, 34:2921-2930.

Heng YHE, McLeay R, Harvey TJ, Smith AG, Barry G, Cato K, Plachez C, Little E, Mason S, Dixon C, Gronostajski RM, Bailey TL, Richards LJ, Piper M (2014) NFIX regulates neural progenitor cell differentiation during hippocampal morphogenesis. Cerebral Cortex, 24; 261-279.

Martynoga B, Mateo JL, Zhou B, Andersen J, Achimastou A, Urban N, van den Berg D, Georgopoulou D, Hadjur S, Wittbrodt J, Ettwiller L, Piper M, Gronostajski RM, Guillemot F (2013) Epigenomic enhancer annotation reveals a key role for NFIX in neural stem cell quiescence. Genes and Development, 27: 1769-1786.

Piper M, Barry G, Hawkins J, Mason S, Lindwall C, Little E, Sarkar A, Smith AG, Moldrich RX, Boyle GM, Tole S, Gronostajski RM, Bailey T, Richards LJ. (2010) NFIA regulates telencephalic progenitor cell differentiation through repression of the Notch effector Hes1. The Journal of Neuroscience, 30(27):9127-9139.

Piper M, Plachez C, Zalucki O, Fothergill T, Guy Goudreau, Erzurumlu R, Gu C, Richards LJ. (2009) Neuropilin 1-Sema signalling regulates crossing of cingulate pioneering axons during development of the corpus callosum. Cerebral Cortex, 19:11-21.

Barry G*, Piper M*, Lindwall C, Moldrich R, Mason S, Little E, Sarkar A, Tole S, Gronostajski RM, Richards LJ (2008) Specific glial populations regulate hippocampal morphogenesis. The Journal of Neuroscience, 28(47): 12328-12340 (* Co-first authors).

Piper M, Dwidevy A, Leung L, Bradley RS, Holt CE (2008) NF-protocadherin and TAF1 regulate axon initiation and elongation in vivo. The Journal of Neuroscience, 28 (1): 100-105.

Piper M, Anderson R, Dwivedy A, Weinl C, van Horck F, Leung KM, Cogill E, Holt C (2006) Signalling mechanisms underlying Slit2-induced collapse of retinal growth cones. Neuron, 49: 215-228. 

Brunet I, Weinl C, Piper M, Trembleau A, Volovitch M, Harris W, Prochiantz A, Holt C (2005) The transcription factor Engrained-2 guides retinal axons.  Nature, 438:  94-98. 

Piper M, Salih S, Weinl C, Holt CE, Harris WA (2005) Endocytosis-dependent desensitization and protein synthesis-dependent resensitization in retinal growth cone adaptation.  Nature Neuroscience, 8:  179-186. 

View all publications from Dr Piper on ORCiD

Group Head



July 2019

Congratulations to James, who has taken a job at the SBMS imaging facility. We look forward to interacting with him productively in coming years!

June 2019

Our manuscript on Usp9x in human patients is now available online.

May 2019

Our paper with the Simmons lab on a novel model for autism is now available online from EBioMedicine.

Well done to Sabrina and Maria, who contributed to a lovely piece of collaborative work with DrLachlan Jolly (University of Adelaide) and Professor Stephen Wood (Griffith University), that was just accepted for publication in Biological Psychiatry.

April 2019

Well done to Cathy Zhang (Simmons lab), for her lovely work on a novel mouse model for autism that has just been accepted in EBioMedicine. We hope this is the first in a long list of collaborative works with the Simmons lab.

Well done to Danyon, who passed his M1. Onwards and upwards!

March 2019

Congrats to Diana, who has been conferred her doctorate from our august institute. Well done!!

Well done to Laura and Sandy, who did very well in the Honours proposal seminars.

February 2019

Congrats to Di, who submitted the corrections for her thesis this month.

Maria sailed through her last milestone presentation; well done you!

Check out the UQ FoM blog featuring Mike discussing Rare Disease Day.

January 2019

Congratulations to Sabrina, whose work on Malan syndrome was recently published in EBioMEdicine. Her work made the cover of this issue!

Welcome also to Laura, who recently joined the lab for her Honours project. Good luck throughout the year Laura.

December 2018

Well done to James, who was awarded the best student poster presentation at the recent EMBL Australia Postgraduate Symposium.

Sonja and Sabrina were both winners at the recent ANS conference. Sonja won best poster, and Sabrina won third prize for the best image competition. Well done also to Sazia and Sabrina for their great talks at ANS.

November 2018

Congratulations to Sabrina, who won the 2018 EMBL Australia Postgraduate Symposium SciArt Competition.

Well done to Raquelle, who received a well-deserved First Class Honours for 2018. Great effort Raquelle!

October 2018

Well done to Lachlan, Sabrina and James, all of whom won prizes at the recent Queensland ANZSCDB meeting, held at the Translational Research Institute. Awesome effort from the Piper lab!

Well done again to Sabrina, who recently won a Poster Award at the 9th SBMS International Post-grad Symposium, for her work on the role of NFIX in macrocephaly

September 2018
Well done to Oressia, as well as Lachlan, Tracey, Danyon and Sabrina, as their paper was recently accepted for publication in the prestigious journal, Cerebral Cortex. An awesome effort!
Well done to Sazia, who was selected to present her work at the recent ComBio conference, that was held in Sydney.

August 2018

Well done to Sazia, who has been selected to give an oral presentation at the 2018 ComBio conference, that will be held in Sydney in September.

Congratulations to Oressia, who recently had a paper published in BMC Neuroscience.

July 2018

Well done to James, who passed his M3 this month.

Sabrina has been selected to participate in the 2018 IMNIS MedTech-Pharma Program. Well done Sabrina!

June 2018

Lachlan and Oressia’s paper on NFIX in the adult hippocampus is now published in Development. Well done all!


Maria was awarded a travel bursary to attend the 2018 ISSCS meeting in Melbourne. Well done Maria!

Mike was awarded the Emerging Leader gong from ANZSCDB for 2018


May 2018

 A big welcome to Dr Sazia Sharmin, who joins us as a UQ Development Fellow this month.

 Well done to Raul, who completed his Summer Scholarship with us this month.

 Congratulations to Lachlan and Oressia, whose paper on identifying cell cycle parameters appeared in the latest issue of The Journal of Molecular Histology


 Mike, and the team from First 5 Forever (State Library of Queensland), collaborated on a video promoting early childhood literacy last year (Little Moments, Big Impact). This video recently won an award at the A corto di libri international film contest.  Well done all!


 April 2018

Welcome to Danyon, who is joining us for a PhD. Danyon will be investigating the role of NFI proteins in adult neurogenesis.

A huge well done to Diana, who submitted her PhD thesis on the role of NFIX in ependymal cell formation this month. An excellent effort, that included two first author manuscripts, and a first author book chapter. Thanks also to Tracey for her contribution to getting Di over the line.

Sabrina waltzed through her M2 this month.  The committee were very impressed!  Well done Sabrina

March 2018

Raquelle did a wonderful job for her Honours proposal talk. Well done Raquelle!

A great effort from Elise and Sabrina, who won bursaries to attend the ISDN meeting in Japan later this year.

Thank you to the Brisbane Marketing and Development Board, who awarded Mike a Trailblazer grant to attend the upcoming ISDN meeting in Nara, Japan. It should be a fantastic conference!

Well done to Elise, who sailed through her M2 this month. The committee were very impressed and excited with the presentation.

Feb 2018

Congratulations to Diana, Raul and Tracey, whose work on the role of NFIX in ependymal cell development was recently published in Neural Development


Exciting news for Elise, Sabrina and Maria, who have all been accepted into the upcoming meeting for the International Society for Developmental Neuroscience, which will be held in Nara (Japan) in May. 


Jan 2018

Another sterling effort by Lachlan, Oressia and Sabrina, whose paper on a method for identifying adult neural stem cells in the hippocampus, was recently published in Developmental Dynamics


Well done to Danyon, who was awarded a First Class Honours for his project last year. Danyon is also joining us for his PhD project.

Welcome to Raquelle Ludwig, who is doing Honours with us this year.

Congratulations to Raul, who got married late last year!

The lab was fortunate to get funding from a number of sources last year; thanks to all those who contributed to this, and to the funding bodies for supporting our research.

We received a Discovery Project Grant from the ARC, with our wonderful collaborator, A/Prof Tom Burne. This work aims to analyse the transcriptional regulation of brain size.

In collaboration with A/Prof Helen Cooper, who was the lead investigator, we also obtained a Project Grant from the NHMRC to investigate the development of hydrocephalus.

We were also fortunate enough to obtain an Explorer Award from the Simons Foundation Autism Research Initiative to study the role of Usp9x in the development of this disorder.



Well done to Elise on the acceptance of her first publication as lead author. Elise's work, detailing the role of NFIX in spinal cord development, was recently accepted to Developmental Biology. Congratulations also to Lachlan and Danyon, who contributed to this work.

Congratulations to Diana, Lachlan and Oressia, who contributed to a paper recently accepted to Brain and Neuroscience Advances. This work was performed in collaboration with the Richards lab at the QBI. Great job all.

Alex recently had a paper accepted for publication in the journal Methods.  This work details her work on the prediction of target genes from transcription factor binding data. Well done Alex!

The lab team, along with wonderful our collaborators Dr. Fiona Simpson (Diamantina Institute), Dr. Glen Boyle (QIMR) and Dr. Ben Panizza (PA Hospital), was fortunate enough be awarded an Enabling Grant from the Australian Skin and Skin Cancer Research Centre for our work in developing an in vitro model for studying perineural invasion of squamous cell carcinomas of the head and neck. We look forward to working with a PhD student in Fiona’s lab, Priscila Oliveira De Lima, who will be driving this work.


Congratulations to James and Sabrina, both of whom were accepted into training courses run through the Neuroscience School of Advanced Studies. Both got to spend a week in Northern Italy learning about the latest techniques in neuroscience. Sabrina also got a travel scholarship from the Graduate School to fund her trip.

Well done to Lachlan and Tracey, who contributed to a recently published study investigating the expression of NFI proteins within the adult brain. Images from the paper made the front cover of the August 1st 2017 issue of The Journal of Comparative Neurology


Elise recently attended the recent EMBL Australia PhD course held at Monash University. Places were scarce for this, so well done Elise!

Congratulations to Lachlan, who recently graduated with a PhD, and who has taken up a prestigious, 4-year post doctoral position at the Crick Institute in London in the lab of Prof. Francois Guillemot. Lachlan’s most recent work on adult neurogenesis was recently published in Developmental Dynamics


James, Alex and Tracey had their work detailing the expression of NFIX within the postnatal cerebellum published recently in the July 2017 Issue of Brain Structure and Function. A job well done all!


Well done to Mitch, who has finished his PhD and has now moved to the States to do a post doc.  Mitch had two recent papers, which appeared in eBioMedicine and The International Journal of Cancer.



Swati had her final piece of work from the lab published in the February 2017 Issue of The Journal of Molecular Histology. Well done to Swati and Tracey on this manuscript!



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