Skip to content Skip to navigation

Team members


Jon Mulholland


Jon Mulholland is Director of the Cell Sciences Imaging Facility (CSIF) at Stanford University. He has served as Director of the CSIF since 2001 during which time he has overseen the expansion of the CSIF by establishing and setting up its current EM core, upgrading significant components of the fluorescence light microscopy core and developing the facility advanced imaging capabilities (super resolution SIM, STORM, gSTED, FLIM, FRET, array tomography). In 2014 he collaborated with the SOE departments of Bioengineering and Chemical Engineering to design, equip and manage a new sister facility in the Shriram Center: The SOE Shriram Center CSIF. Mulholland has extensive training and expertise in light, confocal, 2-photon, super resolution microscopy, FLIM and immunoelectron microscopy. Prior to taking on the role of Director of the CSIF, Mr. Mulholland trained as a cell and molecular biologist while working for 15 years in the laboratory of Dr. David Botstein starting at MIT, then Genentech Inc. and finally Stanford University. While in the Botstein lab he applied fluorescent light and immuno-electron microscopy to study the actin cytoskeleton and membrane vesicle trafficking in yeast. He has collaborated and published with a variety of researchers on a large range of projects. Mr. Mulholland has thirty plus years of experience in both microscopy and laboratory management. Current research interests are centered on establishing large volume 3D EM in the CSIF and correlative light and electron imaging techniques. When not in the facility, he enjoys scuba diving, hiking, and travel.


  • B.S. in Biology/Philosophy, Clark University, MA, USA
  • M.S. in Cell and Molecular Biology, SF State university, CA, USA

Selected Publications

  • Regulation of ER-phagy by a Ypt/Rab GTPase module. (2013). Z. Lipatova, A. H. Shah, J. J. Kim, Jon Mulholland, and Nava Segev. Mol Biol Cell. 2013 Aug 7.
  • Direct interaction between a myosin V motor and the Rab GTPases Ypt31/32 is required for polarized secretion. (2008). Z. Lipatova1, A. A.Tokarev, J. Mulholland, Y Jin, L. S. Weisman, and N. Segev. Mol Biol Cell. 19:4177-87.
  • Immunoelectron Microscopy of Aldehyde-Fixed Yeast Cells. (2002). J. Mulholland and David Botstein. In: Guide to Yeast Genetics and Molecular Biology, 2nd edition, ed. Christine Guthrie and Gerald Fink. Methods in Enzymology
  • Visualization of receptor-mediated endocytosis in yeast. (1999). J. Mulholland, J. Konopka, B. Singer-Kruger, M. Zerial and D. Botstein. Mol. Biol. Cell 10:799-817.
  • The transcriptional program of sporulation in budding yeast. (1998). S. Chu, J. DeRisi, M. Eisen, J. Mulholland, D. Botstein, P. O. Brown, I. Herskowitz. Sci 282(5389):699-705.
  • Yeast actin cytoskeleton mutants accumulate a new class of Golgi-derived secretory vesicles. (1997). J. Mulholland, A. Wesp, H. Riezman, and D. Botstein. Mol. Biol. Cell 8:1481-1499.
  • Ultrastructure of the yeast actin cytoskeleton and Its association with the plasma membrane. J. Mulholland, D. Preuss, A. Moon, A. Wong, D. Drubin and D. Botstein. (1994) J. Cell Biol. 125:381-391.

Cedric Espenel

R&D Scientist Engineer/Shriram Center CSIF Manager

As the manager of the Cell Science Imaging facility (CSIF) at the Shriram Center in Stanford, Cedric Espenel’s primary responsibility is the training of new users in cutting-edge microscopy and imaging techniques and the management of the Shriram Center CSIF's sister facility. Cedric also provides expertise in assembling individual methods into complete image processing pipelines for specific projects using Python/SciPy.

Cedric is a Cell Biologist and Biophysics by doctoral training. In his Ph.D. he investigated the dynamic and partitioning of the tetraspanin CD9 in the plasma membrane using single particle tracking techniques. In addition, he analyzed the dynamics and structure of supported bilayers on different surfaces using atomic force microscopy and fluorescence correlation spectroscopy. As a Postdoctoral Associate, Cedric developed tools to monitor changes in kinesin activity in live cells by FRET-FLIM.


  • B.S. in Cell/Cellular, Molecular Biology and Biochemistry, Montpellier University II, France
  • M.S. in Cell Biology and Biophysics, Montpellier University I&II, France
  • Ph.D. in Cell Biology and Biophysics, Montpellier University I&II, France

Selected Publications

  • Espenel C, Acharya BR, Libanje F, Raingeaud J, Morgan J, Jaulin F, Kreitzer G. KIF17 regulates RhoA-dependent actin remodeling at epithelial cell-cell adhesions. J Cell Sci. 2016.
  • Espenel C, Acharya BR, Kreitzer G. A biosensor of local kinesin activity reveals roles of PKC and EB1 in KIF17 activation. J Cell Biol. 2013 Nov 11.
  • Espenel C, Margeat E, Dosset P, Arduise C, Le Grimellec C, Royer CA, Boucheix C, Rubinstein E, Milhiet PE. Single-molecule analysis of CD9 dynamics and partitioning reveals multiple modes of interaction in the tetraspanin web. J Cell Biol. 2008 Aug 25.
  • Espenel C, Giocondi MC, Seantier B, Dosset P, Milhiet PE, Le Grimellec C. Temperature-dependent imaging of living cells by AFM. Ultramicroscopy. 2008 Sept.


Anum Khan

Advanced Imaging Specialist

As an Advanced Imaging Specialist, Anum overlooks the training of new users in advanced light microscopy techniques including super-resolution microscopy, second harmonic imaging, and FLIM/FRET in addition to confocal microscopy. She is also working alongside the facility Director to set up a CODEX system, in the facility, for multiplex imaging. During her Ph.D., Anum used microscopy-based assays to understand the filament assembly process of cytoskeletal proteins called septins through in vitro reconstitution. Other than microscopy, Anum enjoys hiking, outreach, tv shows and cooking.

Kitty Lee

Light Microscopy Specialist

Kitty has been taking care of the CSIF confocal microscopes and their users since 2002. She's a Hong Kong native who moved to the states in 1994. As a double major in chem and bio with minors in education and computer science, Kitty's college years were spent mostly outside her beloved dorm room (split between cold computer room, creepy bio lab, nasty chem lab, smelly animal facility, busy student health clinic etc, etc). She spent some years at a neurobio lab that specializes in olfactory research, a computational chem lab that seeks the least poisonous cancer-ridding chemical conformation, a cognitive psych lab that tests human response to image stimulus, and a PET scan research that compares human brain images (some of which scored her authorship in a journal publication or two). Her addiction to tantalizing CSIF users stems from her years of tutoring college students and helping clinic nurses handling patients. Latest interest for Kitty outside CSIF includes wilderness first aid class (was once EMT-certified and enjoys playing with fake blood), telemark skiing, snowcamping, kayaking and hiking/backpacking (and shopping for all these activities). Kitty is an avid fencer who enjoys fencing foil and playing with epees, please feel free to challenge her to a duel after your confocal imaging sessions (she has a spare saber as well).


Selected Publications


John Perrino

Electron Microscopy Specialist

John grew up in the suburbs of Cincinnati, Ohio, the 6th of 10 kids. In 1991 he joined Anthony Mahowald’s lab at University of Chicago studying oogenesis and embryogenesis in D. melanogaster. After learning how to prepare fly embryos and ovaries for electron microscopy, John took an in-depth course given by Bob Josephs and Jerry Gross covering sample preparation to image analysis. In July of 1992 he was put in charge of the Cancer Center’s EM facility. John came to Stanford to join the CSIF staff in 2002.


B.S. in Biology, Saint Louis University, MO, USA

Selected Publications

  • Knowland D, Arac A, Sekiguchi KJ, Hsu M, Lutz SE, Perrino J, Steinberg GK, Barres BA, Nimmerjahn A, Agalliu D. Stepwise recruitment of transcellular and paracellular pathways underlies blood-brain barrier breakdown in stroke. Neuron. 2014 May 7.
  • Richardson CE, Spilker KA, Cueva JG, Perrino J, Goodman MB, Shen K. PTRN-1, a microtubule minus end-binding CAMSAP homolog, promotes microtubule function in Caenorhabditis elegans neurons. Elife. 2014 Feb 25.


Ibanri Phanwar-Wood

Array Tomography Specialist

Ibanri joined the CSIF in October 2008. Prior to that, she worked at Stanford’s Department of Comparative Medicine (2002) and the UCSF Medical Center in the Department of Anatomic Pathology (2006). She earned her BS in Zoology and MS in Biochemistry from the North-Eastern Hill University, India.

She has been involved with Electron Microscopy projects in India since 1990 and still assists the EM core at the CSIF as and when needed. Currently she provides service to the Stanford community, out-of-state institutions and out-of-country institutions.

When she is not working, Ibanri enjoys the outdoors, reading, music, movies and spending time with family and friends.


Selected Publications


Marcin Walkiewicz

Scanning Probe Microscopy Laboratory Manager

Marcin is the manager of the scanning probe microscopy laboratory and supervises all shared atomic force microscopes (AFM) and Raman instruments in SNSF as well as the Bruker Bio-AFM in CSIF. He is a factory-trained AFM expert, ready to provide assistance with instrument training, sample preparation, and experiment design. Marcin earned his Ph.D. from University of Virginia and did a postdoctoral fellowship at the National Cancer Institute at the NIH, where he used AFM to study variances in chromatin structure of different human cancer models. Prior to joining SNSF and CSIF he was an Applications Scientist for Bruker Nano Surfaces, where he helped academic and industrial customers find the best solutions to their metrology problems by demonstrating and training them on atomic force microscopes, 3D optical microscopes, and stylus profilers. He also spent extensive time in the field, visiting universities and hosting workshops across North America, where he taught users how to take the full advantage of their instruments.


  • Ph.D. University of Virginia

Lydia-Marie Joubert

Emeritus - Electron Microscopy Specialist

As of September 2017, Lydia is managing her own SEM facility at her alma mater Stellenbosch University in South Africa.

After majoring in Mathematics and Botany for her BS, Lydia attained her PhD in Plant Sciences studying metabolic pathways of monoterpene (essential oils) biosynthesis, using EM autoradiography and microanalytical techniques, and did post-doctoral research on symbiotic endophytic cyanobacteria. Subsequent research on lignocellulolytic microbial enzymes led to an interest in microbial biofilms for bio-energy and bioremediation, and she studied fractal patterns and phenotypic plasticity of complex microbial communities. Visualization with Light and Electron Microscopy has been central to all her research. During her graduate training Lydia was a research fellow at Weizmann Institute, Israel. Lydia joined CSIF in 2006 where she focuses on development of Variable Pressure SEM techniques for imaging non-conductive samples and hydrated biomaterials. In 2009 she received the Professional Technical Staff Award from MSA (Microscopy Society of America) for her Visualization of Hydrogels with Variable Pressure SEM. She is a synchrotron user at SLAC and APS Argonne, applying X-ray Fluorescence to study micronutrients in bacteria. Lydia now focuses on computational 3D-EM techniques through Serial Section Array (SSA)-SEM and Serial BlockFace (SBF)-SEM, while also working on techniques for EM visualization of CLARITY and correlative Light and Electron Microscopy for 3D-EM. Before joining Stanford she earned an MPhil in Higher Education and loves teaching. In 2014 Lydia was a winner in the International Science and Engineering Visualization Challenge (SciVis; Illustration Category), and previously an award winner in Olympus BioScapes digital imaging competition. Her SEM micrographs are included in the Discovery Walk at Stanford Medical School. Lydia’s activities include running, skiing, hiking and Shaolin Kung Fu, while her piano, singing, writing and photography bring joy when at home with her family.


  • B.S. in Botany, Mathematics, Stellenbosch University, South Africa
  • M.S. in Botany, Stellenbosch University, South Africa
  • M.Phil in Higher Education, Stellenbosch University, South Africa
  • Ph.D. in Plant sciences, Indiana University Bloomington, IN, USA

Selected Publications

  • Farnebo, SJ, Woon, CYL, Schmitt, T, Joubert, L-M, Kim, M, Pham, H and Chang, J. 2013. Design and Characterization of an Injectable Tendon Hydrogel: A Scaffold for Guided Tissue Regeneration in the Musculoskeletal System. Tissue Engineering Part A, DOI: 10.1089/ten.TEA.2013.0207
  • Reichelt, M, Joubert, L-M, Perrino, P, Koh, AL, Phanwar, I, Arvin, AM. 2012. Three-dimensional Reconstruction of Herpesvirus Infected Cell Nuclei and PML Nuclear Cages by Serial Section Array Scanning Electron Microscopy and Electron Tomography. PLoS Pathog 8(6): e1002740. doi:10.1371/journal.ppat.1002740.
  • Galic, M., Jeong, S., Tsai, FC, Joubert, L-M, Wu, YI, Hahn, KM, Cui, Y, Meyer, T. 2012. External push and internal pull forces recruit curvature-sensing N-BAR domain proteins to the plasma membrane. Nature Cell Biology 14(8): 874-81 doi: 10.1038/ncb2533.
  • Howitt, MR, Lee, JY, Lertsethtakarn, P, Vogelmann, R, Joubert, L-M, Ottemann, KM, Amieva, MR. 2011. ChePep Controls Helicobacter pylori infection of the Gastric Glands and Chemotaxis in the Epsilonproteobacteria. mBio 2 (4) e00098-11 doi: 10.1128
  • Jiang, X., Malkovskiy, A., Tian, W., Sung, YK, Sun, W, Hau, JL, Manickam, S, Wagh, D, Joubert, LM, Semenza, G, Rajadas, J, Nicolls, MR. 2014. Promotion of airway anastomotic microvascular regeneration and alleviation of airway ischemia by deferoxamine nanoparticles. Biomaterials 35(2), 803-813.
  • Joubert, L-M. 2013. High-resolution imaging with field-emission SEM: the need for downstream plasma cleaning in biological applications. Microscopy and Analysis May: 15-20
  • Joubert, L-M. 2012. VP-SEM: Unsung Hero of SEM Imaging. Application of VP-SEM to beat Visualization challenges. Imaging and Microscopy 4/2012.