Lynd Research Lab: People | Evert K. Holwerda

Evert K. Holwerda

Evert K. Holwerda

Research Scientist
Thayer School of Engineering
Dartmouth College

Group Leader
Fermentation Engineering
Lynd Lab


  • MSc, Bioprocess Technology, Wageningen University and Research 2006
  • PhD, Engineering Sciences, Dartmouth 2013

Research Interests

  • Solubilization and fermentation of lignocellulosic substrates.
  • Physiology of thermophilic anaerobic cellulolytic bacteria.
  • Optimization of data generation, collection and processing.
  • Cultivation techniques for microorganisms, and technologies that increase lignocellulosic substrate utilization.


  1. Yannick J Bomble, Chien-Yuan Lin, Antonella Amore, Hui Wei, Evert K Holwerda, Peter N Ciesielski, Bryon S Donohoe, Stephen R Decker, Lee R Lynd and Michael E Himmel (2017) Lignocellulose deconstruction in the biosphere. Current Opinion in Chemical Biology, 41:61–70

  2. Shuen Hon, Daniel G. Olson, Evert K. Holwerda, Anthony A. Lanahan, Sean J.L. Murphy, Marybeth I. Maloney, Tianyong Zheng, Beth Papanek, Adam M. Guss, Lee R. Lynd. The ethanol pathway from Thermoanaerobacterium saccharolyticum improves ethanol production in Clostridium thermocellum. Metabolic Engineering, Volume 42: 175-184.

  3. Michael L. Balch, Evert K. Holwerda,  Mark F. Davis,  Robert W. Sykes, Renee M. Happs, Rajeev Kumar,  Charles E. Wyman  and Lee R. Lynd (2017). Lignocellulose fermentation and residual solids characterization for senescent switchgrass fermentation by Clostridium thermocellum in the presence and absence of continuous in situ ball-milling. Energy Environmental Science, 10:1252-1261.
  4. Satyakam Dash, Ali Khodayari, Jilai Zhou, Evert K. Holwerda, Daniel G. Olson, Lee R. Lynd and Costas D. Maranas (2017). Development of a core Clostridium thermocellum kinetic metabolic model consistent with multiple genetic perturbations. Biotechnology for Biofuels. 10:108.
  5. Thomas Rydzak, David Garcia, David M. Stevenson, Margaret Sladek, Dawn M. Klingeman, Evert K. Holwerda, Daniel Amador-Noguez, Steven D. Brown, Adam M. Guss (2016). Deletion of Type I glutamine synthetase deregulates nitrogen metabolism and increases ethanol production in Clostridium thermocellum. Metabolic Engineering, Volume 41: 182-191.
  6. Lee R. Lynd, Adam M. Guss, Michael E. Himmel, Dhananjay Beri, Chris Herring, Evert K. Holwerda, Sean J. Murphy, Daniel G. Olson, Julie Paye, Thomas Rydzak, Xiongjun Shao, Liang Tian, and Robert Worthen (2016). Advances in Consolidated Bioprocessing Using Clostridium thermocellum and Thermoanaerobacter saccharolyticum. In: C. Wittman and J.C. Liao (eds.) Industrial Biotechnology: Microorganisms. Wiley-VCH/Verlag.
  7. Tian, L., B. Papanek, D.G. Olson, T. Rydzak, E.K. Holwerda, T. Zheng,  J. Zhou, M. Maloney,  N. Jiang,  R.J. Giannone, R.L. Hettich, A.M. Guss, L.R. Lynd (2016). Simultaneous achievement of high ethanol yield and titer in Clostridium thermocellum. Biotechnology for Biofuels. 9(116).
  8. Beri D., D.G. Olson, E.K. Holwerda, L.R. Lynd (2016). Nicotinamide cofactor ratios in engineered strains of Clostridium thermocellum and Thermoanaerobacterium saccharolyticum. FEMS microbiology letters. 363 (11).
  9. Sand* A., E.K. Holwerda*, N.M. Ruppertsberger, M. Maloney, D.G. Olson, Y. Nataf, I. Borovok, A.L. Sonenshein, E.A. Bayer, R. Lamed, L.R. Lynd and Y. Shoham (2015). Three cellulosomal xylanase genes in Clostridium thermocellum are regulated by both vegetative SigA and alternative SigI6 factors. FEBS letters. 589(20-B):3111-3140.
  10. Holwerda, E.K., P.G. Thorne, D.G. Olson, D. Amador-Noguez, N.L. Engle, T.J. Tschaplinski J.P. van Dijken and L.R. Lynd (2014). The exometabolome of Clostridium thermocellum reveals overflow metabolism at high cellulose loading. Biotechnology for Biofuels. 7(155).
  11. Holwerda, E.K. and L.R. Lynd (2013). Testing alternative kinetic models for utilization of crystalline cellulose (Avicel) by batch cultures of Clostridium thermocellum. Biotechnology and Bioengineering. 110(9):2389-2394.
  12. Holwerda, E.K., L.D. Ellis, L.R. Lynd (2013). Development and Evaluation of Methods to Infer Biosynthesis and Substrate Consumption in Cultures of Cellulolytic Microorganisms. Biotechnology and Bioengineering. 110(9):2380-2388.
  13. Holwerda, E.K., K.D. Hirst and L.R. Lynd (2012). A defined growth medium with very low background carbon for culturing Clostridium thermocellum. Journal of Industrial Microbiology and Biotechnology. 39(6):943-947.
  14. Ellis, L.D., E.K. Holwerda, D. Hogsett, S. Rogers, X. Shao, T. Tschaplinski, P. Thorne and L.R. Lynd (2011). Closing the carbon balance for fermentation by Clostridium thermocellum (ATCC 27405). Bioresource Technology. 103(1):293-299.