Dr. Naoufal Lakhssassi


Last Updated: Feb 01, 2024, 10:08 AM


Plant Breeding/EMS mutagenesis/GWAS/Abiotic Stress/Biotic Stress/ Genetics/Genomics/Epigenetics/Gene editing/Single Cell/Soybean/Tomato


Ph.D. [2011] Universidad de Malaga, Spain: Plant Biology, Plant Pathology, Cell & Molecular Biology

M.S. [2010] Universidad de Malaga, Spain: Biomolecular Engineering and Microbiology

DEA [2007] Universidad de Malaga, Spain: Cell and Molecular Biology

B.S. [2005] University Abdelmalek Essadi, Morocco: Biotechnology


  • Biotechnology

  • Plant genetics

  • Molecular Biology


  • 2022 Inventor of the Year Award. Southern Illinois University System

  • 2018 NSF Conference Award. 17th Biennial Conference Molecular and Cellular Biology of Soybean (Soy 2018).

  • 2017 NSF WSRC10 Conference Award. 10th World Soybean Research Conference

Scholarships / Fellowships:

  • 2012 Post-Doctoral Excellence Fellowship. Universidad de Malaga

  • 2007-2010 PhD Scholarship. Spanish Ministry of Foreign Affairs 

  • 2007 Research Scholarship of Excellence. Junta Committee Andalusia, Spain

  • 2006 DEA Scholarship. Spanish Ministry of Education and Sciences, MEC




  1. Sandeep Goshika, Khalid Meksem, Khaled R Ahmed, and Naoufal Lakhssassi. Deep Learning Model for Classifying and Evaluating Soybean Leaf Disease Damage. International Journal of Molecular Science, 2023.
  2. Nacer Bellaloui, Dounya Knizia, Jiazheng Yuan, Qijian Song, Frances Betts, Teresa Register, Earl Williams, Naoufal Lakhssassi, Hamid Mazouz, Henry T. Nguyen, Khalid Meksem, Alemu Mengistu, and My Abdelmajid Kassem. Genetic Mapping for QTL Associated with Seed Nickel and Molybdenum Accumulation in the Soybean ‘Forrest’ by ‘Williams 82’ RIL Population. Plants, 2023.
  3. Dounya Knizia, Nacer Bellaloui, Jiazheng Yuan, Naoufal Lakhssassi, Erdem Anil, Tri Vuong, Mohamed G. Embaby, Henry T Nguyen, Alemu Mengistu, Khalid Meksem, Abdelmajid My Kassem. QTL and Candidate Genes that Control Seed Sugars Contents in the Soybean ‘Forrest’ By ‘Williams 82’ RIL Population. Plants, 2023.
  4. Sarbottam Piya, Vince Pantalone, Sobhan Bahrami Zadegan, Sarah Shipp, Naoufal Lakhssassi, Dounya Knizia, Hari B Krishnan, Khalid Meksem, Tarek Hewezi. Soybean gene co-expression network analysis identifies two co-regulated gene modules associated with nodule formation and development. Molecular Plant Pathology, 2023.
  5. Naoufal Lakhssassi, Dounya Knizia, Abdelhalim El Baze, Aicha Lakhssassi, Jonas Meksem, and Khalid Meksem. Proteomic, transcriptomic, mutational and functional assays reveal the involvement of both THF and PLP sites at the GmSHMT08 in resistance to soybean cyst nematode. Int. J. Mol. Sci., 2022.
  6. Dounya Knizia, Jiazheng Yuan, Naoufal Lakhssasi, Abdelhalim El Baze, Mallory Cullen, Tri Vuong, Henry T. Nguyen, My Abdelmajid Kassem and Khalid Meksem. QTL and Candidate Genes for Seed Tocopherol Content in ‘Forrest’ By ‘Williams 82’ Recombinant Inbred Line (RIL) Population of Soybean. Plants, 2022.
  7. Naoufal Lakhssassi, Valeria L. Caitar, Dounya Knizia, Mallory A. Cullen, Oussama Badad, Abdelhalim El Baze, Zhou Zhou, Mohamed G. Embaby, Jonas Meksem, Aicha Lakhssassi, Pengyin Chen, Amer AbuGhazaleh, Tri D. Vuong, Henry T. Nguyen, Tarek Hewezi and Khalid Meksem. TILLING-by-Sequencing+ reveals the role of novel Fatty acid desaturases (GmFAD2-2s) in increasing soybean seed oleic acid content. Cells, 2021.
  8. Lakhssassi, N.; Zhou, Z.; Cullen, M.A.; Badad, O.; El Baze, A.; Chetto, O.; Embaby, M.G.; Knizia, D.; Liu, S.; Neves, L.G.; Meksem, K. TILLING-by-Sequencing+ to Decipher Oil Biosynthesis Pathway in Soybeans: A New and Effective Platform for High-Throughput Gene Functional Analysis. Int. J. Mol. Sci. 2021.
  9. Mariola Klepadlo*, Naoufal Lakhssassi*, Gunvant Patil, Tri D. Vuong, Sarbottam Piya, Tarek Hewezi, Robert T. Robbins, Robert M. Stupar, Khalid Meksem, Henry T. Nguyen. Dissecting nematode resistance regions in soybean reveals pleiotropic effect of soybean cyst and reniform nematode resistance genes. Plant Genome, 2021.
  10. Oussama Badad, Naoufal Lakhssassi, Nabil Zaid, Abdelhalim El Baze, Younes Zaid, Jonas Meksem, David A Lightfoot, Huseyin Tombuloglu, El Houcine Zaid, Turgay Unver, Khalid Meksem. High-resolution DNA methylation profiling of wild and cultivated olives reveals Domestication fingerprint on phenolic compounds metabolism. Plants, 2021.
  11. Zhou Zhou, Naoufal Lakhssassi, Dounya Knizia, Abdelhalim El Baze, Zhou Zhou, Cullen Mallory, Oussama Badad, Jonas Meksem, Aicha Lakhssassi, Tri D. Vuong, Henry T. Nguyen, and Khalid Meksem. Characterization and functional genomic studies of soybean acyl-acp thioesterase gene family reveal novel alleles of FAT to alter fatty acid composition in soybean seed. TAG, 2021.
  12. Dounya Knizia, Jiazheng Yuan, Nacer Bellaloui, Tri Vuong, Mariola Usovsky, Qijan Song, Frances Betts, Teresa Register, Earl Williams, Naoufal Lakhssasi, Henry T. Nguyen, Khalid Meksem, Alemu Mengistu, and My Abdelmajid Kassem. The Soybean High Density ‘Forrest’ by ‘Williams 82’ SNP‐Based Genetic Linkage Map Identifies QTL and Candidate Genes for Seed Isoflavone Content. Plants, 2021.
  13. Hassan Hadi Mehdi Al Rubaiy, Ammar Altemimi, Ali Khudair Jaber Al Rikabi, Naoufal Lakhssassi, Anubhav Pratap-Singh. Sustainable Biosynthesis of Antioxidants from Koji Rice Fermented with Aspergillus flavus using Microwave-Assisted Extraction. Applied Sciences, 2021.
  14. Naoufal Lakhssassi, Zhou Zhou, Shiming Liu, Sarbottam Piya, Gunvant B. Patil, Mallory A. Cullen, Abdelhalim El Baz, Oussama Badad, Mohamed G. Embaby, Jonas Meksem, Aicha Lakhssassi, Amer AbuGhazaleh, Michael Fromm, Henry Nguyen, Tarek Hewezi, and Khalid Meksem. Soybean TILLING-by-Sequencing+ reveals the role of novel GmSACPD members in the unsaturated fatty acid biosynthesis while maintaining healthy nodules. Experimental Botany, 2020.
  15. Naoufal Lakhssassi, Azam Baharlouei, Jonas Meksem, Scott D. Hamilton-Brehm, David A. Lightfoot, Khalid Meksem, and Yanna Liang. EMS-induced mutagenesis of Clostridium carboxidivorans for increased atmospheric CO2 reduction efficiency and solvent production. Microorganisms, 2020.
  16. Naoufal Lakhssassi, Sarbottam Piya, Dounya Knizia, Abdelhalim El Baze, Cullen Mallory, Jonas Meksem, Aicha Lakhssassi, Tarek Hewezi, and Khalid Meksem. Mutations at the Serine Hydroxymethyltransferase impact its interaction with a Soluble NSF Attachment Protein and a PathogenesisRelated Protein in soybean. Vaccines, 2020, (8) 349.
  17. Naoufal Lakhssassi, Sarbottam Piya, Sadia Bekal, Shiming Liu, Zhou Zhou, Catherine Bergounioux, Long Miao, Jonas Meksem, Aicha Lakhssassi, Karen Jones, My Abdelmajid Kassem, Moussa Benhamed, Abdelhafid Bendahmane, Kris Lambert, Adnane Boualem, Tarek Hewezi, and Khalid Meksem. A pathogenesis related protein GmPR10-08 promotes a molecular interaction between the GmSHMT08 and GmSNAP18 in resistance to Heterodera glycines. Plant Biotechnology, 2020.
  18. Altemimi, A.B.; Mohammed, M.J.; Yi-Chen, L.; Watson, D.G.; Lakhssassi, N.; Cacciola, F.; Ibrahim, S.A. Optimization of Ultrasonicated Kaempferol Extraction from Ocimum basilicum Using a Box–Behnken Design and Its Densitometric Validation. Foods, 2020, 9, 1379.
  19. Al-sahlany STG, Altemimi AB, Al-Manhel AJA, Niamah AK, Lakhssassi N, and Ibrahim S. Purification of Bioactive Peptide with Antimicrobial Properties Produced by Saccharomyces cerevisiae. Foods, 2020. 9 (3), 324.
  20. Al-Hashimi AG, Ammar AB, Cacciola F, and Lakhssassi N. Development of a Millet Starch Edible Film Containing Clove Essential Oil. Foods, 2020. 9 (2), 184.
  21. Asaad Rehman Saeed Al-Hilphy, Ammar B. Al-Temimi, Hassan Mehdi, Uttpal Anand, Gonzalo Delgado Pando, and Naoufal Lakhssassi. Ultrasound applications in poultry meat processing: A systematic review. Journal of Food Science, 2020. 85(6). DOI: 10.1111/1750-3841.15135.
  22. Alaa R. Abdulstar, Ammar Altemimi, Asaad Rehman Saeed Al-Hilphy, Dennis Watson, and Naoufal Lakhssassi. Water distillation using an Ohmic heating apparatus. International Journal of Ambient Energy, 2020. 10.1080/01430750.2020.1773924.
  23. Jiazheng Yuan, Nacer Bellaloui, Naoufal Lakhssasi, Samer M. AbuBakr, Safiah Kassem, Zakariah Kassem, Sakinah Kassem, Courtney Barnes, Aeisha McLelland, Barbara Brown, Wendy Adams, Tarik El-Mellouki, Khalid Meksem, and My Abdelmajid Kassem. Evaluation of Yield Performance of Soybean Mutant FM6-847 in North Carolina. Atlas Journal of Plant Biology, 2020, pp. 96–105.
  24. Zhou Zhou*, Naoufal Lakhssassi*, Mallory A. Cullen, Abdelhalim El Baz, Tri D. Vuong, Henry T. Nguyen, and Khalid Meksem. Assessment of phenotypic variations and correlation among seed composition traits in mutagenized soybean populations. Genes, 2019. 27;10(12). pii: E975. doi: 10.3390/genes10120975.
  25. Naoufal Lakhssassi, Gunvant Patil, Sarbottam Piya, Zhou Zhou, Azam Baharlouei, Abdelmajid Kassem, David Lightfoot, Dr. Tarek Hewezi, Abdelali Barakat, Henry Nguyen, and Khalid Meksem. Genome reorganization of the GmSHMT gene family in soybean showed a lack of functional redundancy in resistance to soybean cyst nematode. Scientific report, 2019. 9:1506 | https://doi.org/10.1038/s41598-018-37815-w.
  26. Gunvant B. Patil*, Naoufal Lakhssassi*, Jinrong Wan, Li Song, Zhou Zhou, Mariola Klepadlo, Tri D. Vuong, Adrian O. Stec, Sondus S. Kahil, Vincent Colantonio, Babu Valliyodan, Hollis Rice, Sarbottam Piya, Tarek Hewezi, Robert M. Stupar, Khalid Meksem and Henry T. Nguyen. Whole genome re-sequencing reveals the impact of copy number variants of the Rhg4 gene on broad-based resistance to soybean cyst nematode. Plant Biotechnology, 2019. doi: 10.1111/pbi.13086.
  27. Vitor Amorim-Silva, Álvaro García-Moreno, Araceli Castillo, Naoufal Lakhssassi, Alicia Esteban, Abel Rosado, Jéssica Pérez-Sancho, Yansha Li, Pepa Pérez-Rodriguez, Victoriano Valpuesta, Cyril Zipfel, Alberto Macho, Miguel A Botella. TTL proteins scaffold brassinosteroid signaling components at the plasma membrane to optimize signal transduction in Arabidopsis. Plant Cell, 2019. (31):1807–1828, doi/10.1105/tpc.19.00150.
  28. Uttpal Anand, Nadia Jacobo, Ammar Altemimi, and Naoufal Lakhssassi. A Comprehensive Review on Medicinal Plants as Antimicrobial Therapeutics: Potential Avenues of Biocompatible Drug Discovery. Metabolites, 2019. DOI: 10.3390/metabo9110258.
  29. Ammar Altemimi, Salah Naji Aziz, Asaad R. S. Al-HiIphy, Naoufal Lakhssassi, Dennis G. Watson, and Salam A. Ibrahim. Critical review of radio-frequency (RF) heating applications in food processing. Food Quality and Safety, 2019. 1-11. doi:10.1093/fqsafe/fyz002.
  30. Alya Alsaad, Ammar B Altemimi, Salah Naji Aziz, Naoufal Lakhssassi. Extraction and Identification of Cactus Opuntia dillenii Seed Oil and its Added Value for Human Health Benefits. Pharmacognosy Journal, 2019. 11(3):579-587.
  31. James Anderson*, Naoufal Lakhssassi*, Stella K. Kantartzi, and Khalid Meksem. Non-hypothesis Analysis of a Mutagenic Soybean (Glycine max [L.]) Population for Protein and Fatty Acid Composition. Journal of the American Oil Chemists' Society, 2018. DOI 10.1002/aocs.12063.
  32. Lakhssassi N, Colantonio V, Flowers ND, Zhou Z, Henry J, Liu S, Meksem K. Stearoyl-Acyl Carrier Protein Desaturase Mutations Uncover an Impact of Stearic Acid in Leaf and Nodule Structure. Plant Physiology, 2017. Jul;174(3):1531-1543. doi: 10.1104/pp.16.01929. Epub 2017 May 1.
  33. Lakhssassi N, Liu S, Bekal S, Zhou Z, Colantonio V, Lambert K, Barakat A, Meksem K. Characterization of the Soluble NSF Attachment Protein gene family identifies two members involved in additive resistance to a plant pathogen. Scientific Reports, 2017. Mar 24;7:45226. doi: 10.1038/srep45226.
  34. Lakhssassi N, Zhou Z, Liu S, Colantonio V, AbuGhazaleh A, Meksem K. Characterization of the FAD2 Gene Family in Soybean Reveals the Limitations of Gel-Based TILLING in Genes with High Copy Number. Frontiers in Plant Science, 2017. Mar 13;8:324. doi: 10.3389/fpls.2017.00324. eCollection 2017.
  35. Liu S, Kandoth PK, Lakhssassi N, Kang J, Colantonio V, Heinz R, Yeckel G, Zhou Z, Bekal S, Dapprich J, Rotter B, Cianzio S, Mitchum MG, Meksem K. The soybean GmSNAP18 gene underlies two types of resistance to soybean cyst nematode. Nature Communications, 2017. Mar 27;8:14822. doi: 10.1038/ncomms14822.
  36. Kandoth PK, Liu S, Lakhssassi N, Dhroso A, Ludwig AP, Elizabeth KJ, Heinz, Robert H, Amanda P, Wayne LP, Tucke D, Jacobs T, Korkin D, Meksem K, Mitchum M. Role of Rhg4-encoded serine hydroxymethyltransferase in resistance to SCN. Phytopathology, 2017. 107(1): 5.
  37. Kandoth PK, Liu S, Prenger E, Ludwig A, Lakhssassi N, Heinz R, Zhou Z, Howland A, Gunther J, Eidson S, Dhroso A, LaFayette P, Tucker D, Johnson S, Anderson J, Alaswad A, Cianzio SR, Parrott WA, Korkin D, Meksem K, Mitchum MG. Systematic mutagenesis of serine hydroxymethyltransferase reveals essential role in nematode resistance. Plant Physiology, 2017. Nov;175(3):1370-1380. doi: 10.1104/pp.17.00553. Epub 2017 Sep 14.
  38. Meksem K, Liu S, Pramod K, Lakhssassi N, Colantonio V, Kang Jingwen; HR, Yeckel G, Zhou Z, Bekal S, Dapprich, J, Rotter B, Cianzio S, Mitchum, Melissa G. The GmSNAP18 is the Peking-type rhg1-a gene for resistance to soybean cyst nematode. Phytopathology, 2017. 107.
  39. Altemimi A, Lakhssassi N, Baharlouei A, Watson DG, Lightfoot DA. Phytochemicals: Extraction, Isolation, and Identification of Bioactive Compounds from Plant Extracts. Plants, 2017. Sep 22;6(4). pii: E42. doi: 10.3390/plants6040042.
  40. Altemimi A*, Lakhssassi N*, Abu-Ghazaleh A, Lightfoot DA. Evaluation of the antimicrobial activities of ultrasonicated spinach leaf extracts using RAPD markers and electron microscopy. Arch Microbiol., 2017. Dec;199(10):1417-1429. doi: 10.1007/s00203-017-1418-6. Epub 2017 Aug 1.
  41. Al-Saedi SA*, Lakhssassi N*, Kassem MA,  Razaq IB, Ali NA, and  Meksem K. Characterization of diversity of Bradyrhizobia on cowpea in Iraq. Atlas Journal of Biology, 2017. May; pp. 392–401 ISSN 2158-9151.
  42. Bekal S, Domier LL, Gonfa B, Lakhssassi N, Meksem K, Lambert KN. A SNARE-Like Protein and Biotin Are Implicated in Soybean Cyst Nematode Virulence. PLoS One, 2015. Dec 29;10(12):e0145601. doi: 10.1371/journal.pone.0145601. eCollection 2015.
  43. Lakhssassi N, Doblas VG, Rosado A, del Valle AE, Posé D, Jimenez AJ, Castillo AG, Valpuesta V, Borsani O, Botella MA. The arabidopsis TETRATRICOPEPTIDE THIOREDOXIN-LIKE gene family is required for osmotic stress tolerance and male sporogenesis. Plant Physiology, 2012. Mar;158(3):1252-66. doi: 10.1104/pp.111.188920. Epub 2012 Jan 9.

Book Chapter:

  1. Liu Shiming,Naoufal Lakhssassi, Zhou Zhou, Vincent Colantonio, My Abdelmajid Kassem, and Khalid Meksem.Soybean Genomic Libraries, TILLING, and Genetic Resources. Springer-Nature soybean genome book, 2017. DOI https://doi.org/10.1007/978-3-319-64198-0_9.

Patent Applications:

  1. Khalid Meksem, Naoufal Lakhssassi. Soybean Lines with High Seed Protein and Steady to High Oil Content. Publication date. 2023/5/18. Patent office: US. Application number #18050658 https://scholar.google.com/citations?view_op=view_citation&hl=en&user=30-NnXoAAAAJ&cstart=20&pagesize=80&citation_for_view=30-NnXoAAAAJ:_xSYboBqXhAC
  2. Khalid Meksem, Naoufal Lakhssassi, Dounya Knizia, Zhou Zhou. Soybean Lines with Low Saturated Fatty Acid and High Oleic Acid Contents. Publication date: 2022/12/1. Patent office: US. Application number #17827259. https://scholar.google.com/citations?view_op=view_citation&hl=en&user=30-NnXoAAAAJ&cstart=20&pagesize=80&citation_for_view=30-NnXoAAAAJ:D03iK_w7-QYC
  3. Khalid Meksem, Naoufal Lakhssassi, Dounya Knizia. Compositions and Methods to Increase Oleic Acid Content in Soybeans. Publication date 2022/11/17. Patent office: US. Application number #17745148. https://scholar.google.com/citations?view_op=view_citation&hl=en&user=30-NnXoAAAAJ&cstart=20&pagesize=80&citation_for_view=30-NnXoAAAAJ:a0OBvERweLwC
  4. Khalid Meksem, Henry Nguyen, Naoufal Lakhssassi, Mariola Klepadlo, Gunvant Patil. Broad Resistance to Soybean Cyst Nematode. Publication date 2020/8/27. Patent office: US. Application number #16739985. https://scholar.google.com/citations?view_op=view_citation&hl=en&user=30-NnXoAAAAJ&cstart=20&pagesize=80&citation_for_view=30-NnXoAAAAJ:GnPB-g6toBAC

Chairman Workshop Organizer:

  1. Functional Genomics workshop, Plant and Animal Genome Conference, San Diego, CA (since 2020). https://plan.core-apps.com/pag_2023/event/3cee484866e6d00c5e492240c4d5549c.
  2. American Moroccan Agricultural, Health, and Life Sciences (AMAHLS) conference, Morocco (since 2013). https://amahls.org/program/

 Editorial Board Member:

  1. Special issue editor "Plant Development and Hormonal Signaling" in the International Journal of Molecular Sciences (IF= 6.2) since 2023. https://www.mdpi.com/journal/ijms/special_issues/Legume_Genetics
  2. Special issue editor "Advances in Research for Legume Genomics, Genetics, and Breeding 1.0 & 2.0" in the International Journal of Molecular Sciences (IF= 6.2) since 2021. https://www.mdpi.com/journal/ijms/special_issues/_plant_hormonal.
  3. Topical Advisory Panel in Antibiotics journal (IF= 4.63) since 2021. https://www.mdpi.com/journal/antibiotics/topic_editors
  4. Editorial Board Member in SciTechnol journal since 2018. https://www.scitechnol.com/editor-profile/Naoufal_Lakhssassi_PhD/
  5. Board of directors of the High Council of Moroccan American Scholars and Academics (HC-MASA https://hc-masa.org/board-of-directors/