Martins AD

Ana D. Martins, Ph.D.

Post-doctoral Researcher
anacdmartins@gmail.com
ORCID: 0000-0002-7261-9596

Short Biography

Ana D. Martins completed her PhD in Biomedical Sciences at the Institute of Biomedical Sciences Abel Salazar, University of Porto in January 2020. Her LSc and MSc degrees in Biomedical Sciences were completed at the University of Beira Interior in 2011 and 2012, respectively. From 2012 to 2015 she was laboratory technician at the Health Sciences Research Center, University of Beira Interior, Covilhã, Portugal under the supervision of Dr. Pedro F. Oliveira, within the founded project: PTDC/QUI-BIQ/121446/2010. In 2015, she started her PhD under the supervision of Dr. Marco G. Alves (CICS-UBI), and co-supervision of Prof. Mário Sousa (UMIB-ICBAS/UP) and Prof. Rui A. Carvalho (CNC-FCT/UC), with the title: Hormonal regulation of spermatogenesis: role of ghrelin leptin axis and glucagon-like peptide-1 in nutrient membrane transport through Sertoli cell/blood-testis barrier concluded this year.

Currently she is a post-doc fellow within the project: Spermboost - Improving sperm preparation and cryopreservation for Assisted Reproductive Technologies at Institute of Biomedical Sciences Abel Salazar - University of Porto.

Since 2012 she published 39 articles in journals, 4 sections of books, organized 7 events and participated in other 36 events. She participates and/or participated in three founded projects (FCT - PTDC/QUI-BIQ/121446/2010; PTDC/BIM-MET/4712/2014; PTDC/MEC-AND/28691/2017). She had also awarded a Fulbright Fellow in 2017 at Cleveland Clinic, USA under the supervision of Dr. Ashok Agarwal.  

Her research interests are cell hormonal control, cell metabolism and metabolic modulation, particularly in male reproductive system. Her main research keywords are: spermatogenesis; male (in)fertility; Sertoli cell; glucose metabolism; hormonal regulation; metabolic diseases; obesity; metabolism.

Funded projects

PTDC/MEC-AND/28691/2017: “TRAITor - Tracking TRAnsgenerational epigenetic Inheritance of metabolic diseases Transmitted via overweight or obese fathers”. P.I.: Marco G Alves

PTDC/BIM-MET/4712/2014: “Obesitility – Gut hormones and testicular metabolism: mechanisms associated with obesity-related male infertility”. P.I.: Marco G Alves

Spermboost: improving sperm preparation and cryopreservation for Assisted Reproductive Technologies”, P.I.: Marco G Alves by Merck®

Ended funded projects

PTDC/QUI-BIQ/121446/2010:“Can Hormonal (De)regulation of Ion Transporters in Human Sertoli Cells be Responsible for Infertility?” - P.I.: Pedro Fontes Oliveira.

Fellowships and Prizes

20222 - Stimulus for Scientific Employment with the project: "No Fertility for Old Men - Identification of spermatozoa and seminal plasma biomarkers for capacitation in aging using a proteomic approach"

2020 - Post-Doctoral fellow in the project “Spermboost: Improving Sperm Preparation and Cryopreservation for Assisted Reproductive Technologies”

2019 - Travel grant to attend the Molecular Andrology 2019 symposium at the Justus Liebig University, Giessen, Germany, 24-26 September 2019.

2018 - Travel grant to attend the ICDM 2018, held from 11-13 October, at the Grand Hilton Hotel Seoul, Seoul, South Korea.

2018 - Fellowship to participate at the MiPschool Tromsø-Bergen, 20-24 October, Hurtigruten: Tromsø - Bergen, Norway. 

2017 - Fellowship to participate at the 17th FEBS Young Scientists’ Forum and at the 42nd FEBS congress. 7-14 September 2017, Jerusalem, Israel.

2017 - Fulbright research scholarship with the support from the Portuguese Foundation for Science and Technology (FCT) for the academic year 2017/2018.

2016 - Fellowship to participate in the 2nd International Workshop on Klinefelter Syndrome. 10-12 March, 2016, Munster, Germany.

2015 - 4-year PhD fellowship from the Portuguese Foundation for Science and Technology (FCT).

2015 - Fellowship from the ISMAR - International Society of Magnetic Resonance to participate in the ISMAR 2015 Conference. August 16-21, Shanghai, China

2015 - Fellowship to participate in the 40th Congress of The Federation of European Biochemical Societies (FEBS). July 4-9, Berlin, Germany

2012 - Research fellow in the project: “Can Hormonal (De)regulation of Ion Transporters in Human Sertoli Cells be Responsible for Infertility?”

Proceedings in Peer-reviewed Journals

17. AD Martins, Monteiro MP, Sousa M, Barros A, Silva J, Carvalho RA, Silva BM, Oliveira PF, Alves MG (2016) "Glucagon-like peptide-1 stimulates lactate production by human Sertoli cells with possible implication to male reproductive potential". 50th Annual Scientific Meeting of the European Society for Clinical Investigation in European Journal of Clinical Investigation. 46(S1):120. ISSN: 1365-2362

16. Silva BM, Tomas GD, Dias TR, Martins AD, Rato L, Alves MG, Oliveira PF (2016). Sperm quality of pre-diabetic wistar rats restored after a diet with white tea. Medicine. 95(10): e02371 (DOI:10.1097/MD.0000000000002371)

15. Almeida SP, Martins AD, Jarak I, Barros A, Silva J, Sousa M, Oliveira PF, Alves MG (2015) Testicular metabolic biomarkers in men with Klinefelter syndrome. Medicine. 95(10): e02371 (DOI:10.1097/MD.0000000000002371)

14. Martins AD, Morreira AC, Sá R, Monteiro MP, Sousa M, Carvalho RA, Silva BM, Oliveira PF, Alves MG (2015). Leptin modulates human Sertoli cells glucose metabolism: relevance for obesity-induced male infertility. Medicine. 95(10): e02371 (DOI:10.1097/MD.0000000000002371)

13. Martins AD, Moreira AC, Sá R, Monteiro MP, Sousa M, Carvalho RA, Silva BM, Oliveira PF, Alves MG (2016). Ação da leptina no metabolismo da glucose das células de Sertoli: impacto na fertilidade masculina. CAPTAR. 6(1): 43 – 44. ISSN:1647-323X

12. Jesus TT, Bernardino RL, Martins AD, Sá R, Sousa M, Alves MG, Oliveira PF (2016) As aquaporinas-4 e -9 são expressas em células de Sertoli de rato e interagem com o CFTR. CAPTAR. 6(1):38-9. ISSN:1647-323X

11. Dias TR, Alves MG, Bernardino RL, Martins AD, Moreira AC, Silva J, Barros A, Sousa M, Silva BM, Oliveira PF (2016) Caffeine and male fertility: Metabolic effects and oxidative profile in human Sertoli cells. CAPTAR. 6(1): 28 – 29. ISSN:1647-323X

10. Bernardino RL, Martins AD, Costa AR, Silva J, Barros A, Sousa M, Alves MG, Oliveira PF (2016) Influência do estradiol nos transportadores membranares de bicarbonato dependentes de sódio da família SLC4 em células de Sertoli humanas. CAPTAR. 6(1): 4 – 6. ISSN:1647-323X

9. Pinto S, Martins AD, Moreira PI, Carvalho RA, Sousa M, Barros A, Silva J, Simões T, Oliveira PF, Alves MG (2015) Metabolic fingerprints in testicular biopsies from type 1 diabetic patients. Hum Reprod 30(Suppl. 1):170.

8. Alves MG, Bernardino RL, Martins AD., Jesus TT, Sousa M, Sá R, Oliveira PF (2015) Estrogenic regulation of bicarbonate transporters from SLC4 family in rat Sertoli cells. Human Reproduction. 30(Suppl. 1):161.

7. Martins AD, Sá R, Monteiro MP, Barros A, Silva J, Sousa M, Carvalho RA, Silva BM, Oliveira PF, Alves MG (2015). Ghrelin modulates human Sertoli cells metabolism: relevance for male fertility. FEBS J. 282 (Suppl. 1) 55: 188. 

6. Bernardino RL, Martins AD, Costa AR, Silva J, Barros A, Sá R, Sousa M, Alves MG, Oliveira PF (2015). Estrogenic regulation of Na+-dependent bicarbonate transporters from SLC4 family in human Sertoli Cells. FEBS J. 282 (Suppl. 1) 55: 199.

5. Dias TR, Alves MG, Bernardino RL, Martins AD, Moreira AC, Silva J, Barros A, M. Sousa, Silva BM, Oliveira PF (2015). Caffeine-induced alterations in human Sertoli cells metabolism and oxidative profile. FEBS J. 282 (Suppl. 1) 55: 386.

4. Correia S, Alves MR, Martins AD, Cavaco JE, Oliveira PF, Socorro S (2014) Effect of 17b-estradiol on germ cell apoptosis by modulating the expression of SCF/c-kit, FasL/FasR and regucalcin. Endocrine Abstracts 35: P702. (DOI: 10.1530/endoabs.35.P702).

3. Alves MG*, Jesus TT*, Bernardino RL, Martins AD, Sá R, Sousa M, Oliveira PF (2013) Water transport in seminiferous tubule: aquaporins identification in Sertoli cells. Proc. 37th IUPS, LB91

2. Bernardino RL, Martins AD, Jesus TT, Sá R, Sousa M, Socorro S, Alves MG, Oliveira PF (2013) Regulation by estrogens of bicarbonate secretion mechanisms in Sertoli cell. Proc. 37th IUPS, LB113. 

1. Martins AD, Simões VL, Alves MG, Cavaco JE, Socorro S, Oliveira PF (2012) Hormonal Regulation of Na+­H+ Exchanger 3 (NHE­3) in Sertoli Cells: A Possible Role for Estrogens in Spermatogenesis. J. Reproduktionsmed. Endokrinol. 9(5): 371.

Publications

50. Moreira R, Martins AD, Ferreira R, Alves MG, Pereira MdL, Oliveira PF. Impact of Chromium Picolinate on Leydig Cell Steroidogenesis and Antioxidant Balance Using an In Vitro Insulin Resistance Model. Antioxidants. 2024; 13(1):40. https://doi.org/10.3390/antiox13010040

49. Martins AD, Ribeiro JC, Ferreira R, Alves MG, Oliveira PF. Understanding the age-related alterations in the testis-specific proteome, Expert Review of Proteomics, 2023 DOI: 10.1080/14789450.2023.2274857

48. Mateus FG, Moreira S, Martins AD, Oliveira PF, Alves MG, Pereira MdL. L-Carnitine and Male Fertility: Is Supplementation Beneficial? Journal of Clinical Medicine. 2023; 12(18):5796. https://doi.org/10.3390/jcm12185796

47. Moreira R, Martins AD, Alves MG, de Lourdes Pereira M, Oliveira PF. A Comprehensive Review of the Impact of Chromium Picolinate on Testicular Steroidogenesis and Antioxidant Balance. Antioxidants. 2023; 12(8):1572. https://doi.org/10.3390/antiox12081572

46. Correia AS, Pereira SC, Morais T, Martins AD, Monteiro MP, Alves MG, Oliveira PF. Obesity-Related Genes Expression in Testes and Sperm Parameters Respond to GLP-1 and Caloric Restriction. Biomedicines. 2022 Oct 17;10(10):2609. doi: 10.3390/biomedicines10102609

45. Silva JV, Santiago J, Matos B, Henriques MC, Patrício D, Martins AD, Duarte JA, Ferreira R, Alves MG, Oliveira P, Oliveira PF, Fardilha M. Effects of Age and Lifelong Moderate-Intensity Exercise Training on Rats' Testicular Function. Int J Mol Sci. 2022 Oct 1;23(19):11619. doi: 10.3390/ijms231911619
43. Ribeiro, J.C.; Martins, A.D.; Jarak, I.; Carvalho, R.A.; Alves, M.G.; Oliveira, P.F. (2022) Exenatide and Dapagliflozin Combination Enhances Sertoli Cell Secretion of Key Metabolites for Spermatogenesis. Biomedicines, 10, 1115. doi: 10.3390/biomedicines10051115

42. Ribeiro JC, Braga PC, Martins AD, Silva BM, Alves MG, Oliveira PF (2021). Antioxidants Present in Reproductive Tract Fluids and Their Relevance for Fertility. Antioxidants, 10(9), 1441. doi:10.3390/antiox10091441

41. Pereira SC, Martins AD, Monteiro MP, Pinto S, Barros A, Oliveira PF, Alves MG (2021). Expression of obesity-related genes in human spermatozoa affects the outcomes of reproductive treatments. F&S Science. doi:10.1016/j.xfss.2021.03.004 

40. Pereira SC, Martins AC, Moreira BP, Bernardino RL, Barros A, Monteiro MP, Oliveira PF, Alves MG (2021) Obesity-related genes are expressed in human Sertoli cells and modulated by energy homeostasis regulating hormones. Journal of Cellular Physiology, . doi:10.1002/jcp.30230

39. Moreira BP, Silva AM, Martins AD, Monteiro MP, Sousa M, Oliveira PF, Alves MG (2020) Effect of Leptin in Human Sertoli Cells Mitochondrial Physiology. Reproductive Sciences, 10.1007/s43032-020-00328-x

38. Martins AD, Selvam MKP, Agarwal A, Alves MG, Baskaran S. (2020) Alterations in seminal plasma proteomic profile in men with primary and secondary infertility. Scientific Reports. 10, 7539. DOI: 10.1038/s41598-020-64434-1

37. Martins AD, Jarak I, Morais T, CarvalhoRA, Oliveira PF, Monteiro MP, Alves MG (2020) Caloric restriction alters thehormonal profile and testicular metabolome resulting in alterations of spermhead morphology. Am J Physiol Endocrinol Metab. 318(1): E33-E43.  DOI:10.1152/ajpendo.00355.2019

36. Martins AD, Oliveira PF, Alves MG(2019) Assessment of Sertoli cell proliferation by3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyltetrazolium bromide and sulforhodamineB assays. Curr. Protoc. Toxicol., 81, e85. DOI: 10.1002/cptx.85

35. Martins AD, Monteiro MP, Silva BM,Barros A, Sousa M, Carvalho RA, Oliveira PF, Alves MG (2019) Metabolic dynamicsof human Sertoli cells are differentially modulated by physiological andpharmacological concentrations of GLP-1. Toxicol. Appl. Pharmacol. 362:1-8. DOI:10.1016/j.taap.2018.10.009

34. Martins AD, Agarwal A, Baskaran S,Pushparaj NP, Ahmad G, Selvam MPK (2019) Alterations of Spermatozoa ProteomicProfile in Men with Hodgkin's Disease Prior to Cancer Therapy. World J. MensHealth. 2019;37: e32 DOI: 10.5534/wjmh.190012

33. Selvam MKP, Agarwal A, Dias TR, MartinsAD, Baskaran S, Samanta L (2019) Round cells do not contaminate or mask humansperm proteome in proteomic studies using cryopreserved samples. Andrologia.2019 Jun 5: e13325. DOI: 10.1111/and.13325

32. Selvam MKP, Agarwal A, Dias TR, MartinsAD, Samanta L (2019) Presence of round cells proteins do not interfere withidentification of human sperm proteins from frozen semen samples by LC-MS/MS.‎Int. J. Mol. Sci., Jan 14;20(2). pii: E314.  DOI: 10.3390/ijms20020314

31. Selvam MKP, Agarwal A, Sharma R,Samanta L, Gupta S, Dias TR, Martins AD (2019) Protein Fingerprinting ofSeminal Plasma Reveals Dysregulation of Exosome-Associated Proteins inInfertile Men with Unilateral Varicocele. World J. Mens Health. 2019 Apr 3.DOI: 10.5534/wjmh.180108

30. Martins AD, Agarwal A (2019). Oxidation reduction potential: a new biomarker of male infertility. Panminerva Medica, Jun;61(2):108-117. DOI:10.23736/S0031-0808.18.03529-2

29. Selvam MKP, AgarwalA, Dias TR, Martins AD, Baskaran S, Samanta L. (2018). Molecularpathways associated with sperm biofunction are not affected by the presence ofround cells and leukocytes proteins in human sperm proteome. J. Proteome Res.,18(3):1191-1197. DOI: 10.1021/acs.jproteome.8b00829

28. Martins AD, Majzoub A, Agarwal A (2018) Metabolic Syndrome and Male Fertility. World J. Mens Health. 36: e37. DOI:10.5534/wjmh.180055

27. Martins AD, Sá R,Monteiro MP, Barros A, Sousa M, Carvalho RA, Silva BM, Oliveira PF, Alves MG(2016). Ghrelin dysfunction may alter the reproductivepotential of males by disrupting the nutritional support ofspermatogenesis.  Act. Farm. Port.5(2):121-127. 

26. Martins AD, Sá R, Monteiro MP, Barros A, Carvalho RA, Silva BM, Oliveira, PF, Alves MG (2016). Ghrelin acts as energy status sensor of male reproduction by modulating Sertoli cells glycolytic metabolism and mitochondrial bioenergetics. Mol. Cell. Endocrinol. 434:199-209 (DOI: 10.1016/j.mce.2016.07.008)

25. Bernardino RL, Costa AR, Martins AD, Silva J, Barros A, Sousa M, Sá R, Alves MG, Oliveira PF (2016) Estradiol modulates Na+ -dependent HCO3 - transporters altering intracellular pH and ion transport in human Sertoli cells: A role on male fertility? Biol Cell. 108(7): 179-188 (DOI: 10.1111/boc.201500094)

24. Alves MG, Martins AD, Jarak I, Barros A, Silva J, Sousa M, Oliveira PF (2016) Testicular lactate content is compromised in men with Klinefelter syndrome. Mol. Reprod. Dev. 83(3):208-216 (DOI: 10.1002/mrd.22608) 

23. Oliveira PF*, Tomás GD*, Dias TR, Martins AD, Rato L, Alves MG, Silva BM (2015) White tea consumption restores sperm quality in prediabetic rats preventing testicular oxidative damage. Reprod Biomed Online. 31(4): 544-556 (DOI: 10.1016/j.rbmo.2015.06.021)

22. Bernardino RL, Martins AD, Jesus TT, Sá R, Sousa M, Alves MG, Oliveira PF (2015) Estrogenic regulation of bicarbonate transporters from SLC4 family in rat Sertoli Cells. Molecular and Cellular Biochemistry. 408(1-2):47-54 (DOI:10.1007/s11010-015-2481-4).

21. Martins AD, Moreira AC, Sá R, Monteiro MP, Sousa M, Carvalho RA, Silva BM, Oliveira PF, Alves MG. Leptin modulates human Sertoli cells acetate production and glycolytic profile: a novel mechanism of obesity-induced male infertility? BBA-Mol Basis of Disease 1852(9): 1824-1832 (DOI:10.1016/j.bbadis.2015.06.005)

20. Alves MG, Martins AD, Moreira PI, Carvalho RA, Sousa M, Barros A, Silva J, Pinto S, Simões T, Oliveira PF. Metabolic fingerprints in testicular biopsies from type 1 diabetic patients. Cell Tiss Res. (DOI:10.1007/s00441-015-2217-5)

19. Alves MG, Martins AD, Teixeira NF, Rato L, Oliveira PF, Silva BM (2015) White tea consumption improves cardiac glycolytic and oxidative profile of prediabetic rats. Journal of Functional Foods. 14:102-110 (DOI:10.1016/j.jff.2015.01.019)

18. Dias TR, Alves MG, Bernardino RL, Martins AD, Moreira AC, Silva J, Barros A, Sousa M, Silva BM, Oliveira PF (2015) Dose-dependent effects of caffeine in human Sertoli cells metabolism and oxidative profile. Toxicology. 328:12-20 (DOI:10.1016/j.tox.2014.12.003)

17. Oliveira PF, Martins AD, Moreira AC, Cheng CY, Alves MG (2015) The Warburg effect revisited - lesson from the testis. Med. Res. Rev. 35(1):126-51 (DOI: 10.1002/med.21325)

16. Rocha CS, Rato L, Martins AD, Alves MG, Oliveira PF (2015) Melatonin and male reproductive health: relevance of darkness and antioxidant properties. Curr. Mol. Med. 15(4): 299-311. (DOI:10.2174/1566524015666150505155530)

15. Jesus TT, Bernardino RL, Martins AD, Sá R, Sousa M, Alves MG, Oliveira PF (2014) Aquaporin-9 is expressed in rat Sertoli cells and interacts with the cystic fibrosis transmembrane conductance regulator. IUBMB Life 66(9): 639–644 (DOI 10.1002/iub.1312)

14. Rocha CS, Martins AD, Rato L, Silva BM, Oliveira PF, Alves MG (2014) Melatonin alters the glycolytic profile of Sertoli cells: implications for male fertility. Mol. Hum. Reprod. 20(11):1067-76 (DOI:10.1093/molehr/gau080)

13. Martins AD*, Bernardino RL*, Neuhaus-Oliveira A, Sousa M, Sá R, Alves MG, Oliveira PF (2014) Physiology of Na+/H+ Exchangers in the Male Reproductive Tract: Relevance for Male Fertility. Biol. Rep. 91(1):11. (DOI: 10.1095/​biolreprod.114.118331)

12. Jesus TT, Martins AD, Bernardino RL, Sá R, Sousa M, Alves MG, Oliveira PF (2014) Aquaporin-4 as a molecular partner of cystic fibrosis transmembrane conductance regulator in rat Sertoli cells. Biochem. Biophys. Res. Com. 446(4): 1017–1021 (DOI: 10.1016/j.bbrc.2014.03.046).

11. Oliveira PF, Alves MG, Martins AD, Correia S, Bernardino RL, Silva J, Barros A, Sousa M, Cavaco JE, Socorro S. (2014) Expression pattern of G protein-coupled receptor 30 in human seminiferous tubular cells. Gen. Comp. Endoc. 201:16-20 (DOI: 10.1016/j.ygcen.2014.02.022).

10. Martins AD, Alves MG, Bernardino RL, Dias TR, Silva BM, Oliveira PF. (2014) Effect of white tea (Camellia sinensis (L.)) extract in the glycolytic profile of Sertoli cell. Eur. J. Nut. (DOI: 10.1007/s00394-013-0640-5).

9. Martins AD*, Alves MG*, Vaz CV, Correia S, Moreira PI, Oliveira PF, Socorro S. (2014) Metformin and male reproduction: effects in Sertoli cell metabolism. British J. Pharm. 171(4): 1033–1042.

8. Dias TR, Martins AD, Reis VP, Socorro S, Silva BM, Alves MG, Oliveira PF (2013) Glucose transport and metabolism in Sertoli cell: relevance for male fertility. Cur. Chem. Biol. 7(3):282-93. (DOI: 10.2174/2212796807999131128125510)

7. Bernardino RL, Martins AD, Socorro S, Alves MG, Oliveira PF (2013) Effect of pre-diabetes on membrane bicarbonate transporters in testis and epididymis. J. Memb. Biol. 246(12):877-83 (DOI:10.1007/s00232-013-9601-4)

6. Dias TR, Rato L, Martins AD, Simões AD, Jesus TT, Alves MG, Oliveira PF (2013) Insulin deprivation decreases caspase-dependent apoptotic signaling in cultured rat Sertoli cells. ISRN Urology ID:970370. (DOI:10.1155/2013/970370)

5. Martins AD*, Alves MG*, Simões VL, Dias TD, Rato L, Moreira PI, Socorro S, Cavaco JE, Oliveira PF (2013) Control of Sertoli cell metabolism by sex steroid hormones is mediated through modulation in glycolysis-related transporters and enzymes. Cell Tissue Res. 354(3):861-8. (DOI:10.1007/s00441-013-1722-7)

4. Bernardino RL*, Jesus TT*, Martins AD, Sousa M, Barros A, Cavaco JE, Socorro S, Alves MG, Oliveira PF (2013) Molecular basis of bicarbonate membrane transport in the male reproductive tract. Curr. Med. Chem. 20(32):4037-49. (DOI: 10.2174/15672050113109990200)

3. Alves MG, Martins, AD, Socorro S, Cavaco JE, Oliveira PF (2013) Diabetes, insulin-mediated glucose metabolism and Sertoli/blood-testis barrier function. Tissue Barriers. 1(2):1-12. (DOI:10.4161/tisb.23992)

2. Alves MG, Martins AD, Rato L, Moreira PI, Socorro S, Oliveira PF (2013) Molecular mechanisms beyond glucose transport in Diabetes-related male infertility. Bioch. Biophys. Acta. 1832(5): 626-35. (DOI:10.1016/j.bbadis.2013.01.011)

1. Simões VL, Alves MG, Martins AD, Dias TD, Rato L, Socorro S, Oliveira PF (2013) Regulation of Apoptotic Signalling Pathways by 5α-dihydrotestosterone and 17β-estradiol in Immature Rat Sertoli Cells. J. Steroid Biochem. Mol. Biol. 135(1): 15-23. (DOI:10.1016/j.jsbmb.2012.11.019)


* both authors contributed equally