First name Last Name Year(s) Designator Home Country (from) Title Research PhD Institution Thesis
Monica Santos-Mendoza 2011 Borlaug Fellow Mexico Transcriptomic and Motobolomic of Cocoa embriogenesis
Leidy Paola Delgadillo Duran 2020 Borlaug Fellow Colombia Cd/ CAS9
Peter Aikpokpodion 2006-2007 Borlaug Fellow Nigeria Genetic Diversity in Cacao (Theobroma cacao L.) Collections in Nigeria Determined by Phenotypic and Simple Sequence Repeat (SSR's) Markers
Chia-Lock Tan 2006-2007 Fulbright Scholar Malasia Molecular and Biotechnological Techniques for the Improvement of Cocoa Against Pests
Antonio Figueira 2007-2008 Fulbright Scholar Brazil Global Analysis of Genes Differentially Expressed in the Theobroma Cacao Genotypes in Response to Witches Broom Infection
Nicolas Dr. Niemenak 2010-2011 Fulbright Scholar Cameroon Maturation and germination physiology of cacao
Catalina Chapparo 2001 Visiting Scientist Colombia Cacao Somatic Embryogenesis for Colombian Varieties
Anna Muyiwa 2008 Visiting Scientist Nigeria Cacao, germplasm & germination
Yaleidis Mendez 2010 Visiting Scientist Puerto Rico Cacao somatic embryogenesis
Claudia Arevalo 2012 Visiting Scientist Venezuela Cacao Transformation
Dadzie Abu 2013 Visiting Scientist Ghana Disease resistance in cacao
Suzana Sato Guima 2015 Visiting Scientist Brazil Knowledge sharing
Lin Zhou 2015 Visiting Scientist China Defense gene in diversity population
Adrielle de Vasconcelos 2015 Visiting Scientist Brazil Knowledge sharing
Annelle Holder-John 2016 Visiting Scientist Trinidad & Tobago Cutting evaluation in two species of cacao
Jaime Osorio 2016 Visiting Scientist Colombia Cadium uptake in cacao
Gabriela Silva-Pumarada 2018 Visiting Scientist Puerto Rico Cacao
Fernando Santiago 2018 Visiting Scientist Puerto Rico Cacao
Thierry Legrave 2019 Visiting Scientist France CRISPR disease genes
Rosalia Velasquez-Salazar 2008-2009 Visiting Scientist Venezuela Cacao Somatic Embryogenesis/ Jatropha Somatic Embryogenesis
Luis Mejia-Franco 2009, 2012 Visiting Scientist Panama Cacao Endophytes
Desire Pokou 2013, 2016, 2017 Visiting Scientist Ivory Coast Disease resistance in cacao
Marieke Hilgers 2019-2020 Visiting Scientist Netherlands Functional genomics
Adriana Gallego 2106-2017 Visiting Scientist Colombia Polyphenol light effect
Siela Maximova 1996 Post Doctoral Scholar Research Assistant Cacao Ph.D. Penn State
Yuan Yao 2002 Post Doctoral Scholar Research Assistant Maize Ph.D. Jiangnan University, China
Lauren Gross 1992-1993 Post Doctoral Scholar Research Assistant Plant Physiology Ph.D. Penn State
Nancy Eckardt 1992-1993 Post Doctoral Scholar Research Assistant Maize Ph.D. Penn State
Leslie Carlini 1994-1997 Post Doctoral Scholar Research Assistant Plant Physiology Ph.D. Penn State
Zhijian Li 1996-1998 Post Doctoral Scholar Research Assistant Cacao Ph.D. Louisiana State University
Susan Blauth 1997-1999 Post Doctoral Scholar Research Assistant Maize Ph.D. Cornell University
Joe Verica 2002-2007 Post Doctoral Scholar Research Assistant Cacao Ph.D. Penn State
Jihong Li 2004-2005 Post Doctoral Scholar Research Assistant Maize Ph.D. Hainan University, China
Marna Yandeau-Nelson 2005-2008 Post Doctoral Scholar Research Assistant Maize Ph.D. Iowa State University
Luis Mejia-Franco 2009-2010 Post Doctoral Scholar Research Assistant Tree Symbiont Interactions Ph.D. Rutgers University
Zi Shi 2010-2012 Post Doctoral Scholar Research Assistant Bread Project/ Pathogense Ph.D. Penn State
Xiangling Shen 2011-2012 Post Doctoral Scholar Research Assistant Cocoa propagation/ microarray analysis and bioinformatics Ph.D. Nanjing Agricultural University, China
Emily Helliwell 2013-2014 Post Doctoral Scholar Research Assistant Cacao Ph.D. Penn State
Yufan Zhang 2015-2016 Post Doctoral Scholar Research Assistant Cacao Ph.D. Penn State
Andrew Fister 2016-2018 Post Doctoral Scholar Research Assistant Cacao Ph.D. Penn State
Akiva Shalit-Kaneh 2017-2019 Post Doctoral Scholar Research Assistant Cacao Ph.D. University of California, Davis
Sarah Prewitt 2017-2020 Post Doctoral Scholar Research Assistant Cacao Ph. D. University of North Texas
Patrick Thomas 2023-2025 Post Doctoral Scholar Research Assistant Cacao Ph.D. University of California, Riverside
Jigeesha Mukhopadhyay 2024-2025 Post Doctoral Scholar Research Assistant Cacao Ph.D. University of Manitoba, Canada
Mark Guiltinan 1986 Graduate Student Ph.D. Biology University of California, Irvine Thesis: The Isolation, Characterization and Intergeneric Transfer of Two Soybean (Glycine Max L.) Beta-tubulin Genes (TI Plasmid)
Charlene Chang 1993 Graduate Student M.S. Plant Physiology Penn State Thesis: Optimization of in vitro regeneration of geranium (Pelargonium xhortorum L.H. Bailey) for development of a genetic transformation system.
Dane Fisher 1995 Graduate Student Ph.D. in Genetics Penn State Thesis: Molecular genetic analysis of multiple isoforms of starch branching enzyme with emphasis on Zea mays L.; Past Position: Pfeiffer University, NC (2010)
Xiping Niu 1995 Graduate Student Ph.D. in Plant Physiology Penn State Thesis: DNA binding specificity and interactions with nucleosomal DNA of the plant bZip protein EmBP-1; Past Position: Pionneer Hi-Bred International (2010)
Sashi Prabhakar 1995 Graduate Student M.S. Plant Physiology Penn State Thesis: cDNA cloning and characterization of a EmBP-1 homologue in maize (Zea mays): A basic leucine zipper transcription factor; Past Position: Sr. Analyst Application Svcs, Merck Research Labs
Ming Gao 1996 Graduate Student Ph.D. in Plant Physiology Penn State Thesis: Molecular characterization of starch branching enzyme genes, SbeI, Sbe2b and Sbe2a in maize (Zea mays L.); Past Position: Plant Biiotechnology Institute, NRC, Saskatoon (2010)
Siela Maximova 1996 Graduate Student Ph.D. in Plant Physiology Penn State Thesis: Development of a genetic transformation system for apple; Past Position: Senior Scientist and Professor of Horticulture, Penn State University, Dept. of Plant Science. Current email: snm104@psu.edu
Kyung-Nam Kim 1997 Graduate Student Ph.D. in Plant Physiology Penn State Thesis: Isolation and promoter analysis of the genes for starch branching enzyme in maize 2010; Past Position: Associate Professor, Dept. Of Molecular Biology, Sejong University; knkim@sejong.ac.kr
Abdoulaye Traore 1999 Graduate Student Ph.D. in Food Science Penn State Thesis: Tissue culture and Transformation of cacao; Past Position: Process & Product Development Scientist; Mars Chocolate US-ELZ T: 717-367-0991; M: 717-538-4983; Fax: 717-361-4608 (2010); Email: abdoulaye.traore@effem.com
Jennifer Tomscha 2001 Graduate Student Ph.D. Horticulture Penn State Thesis: Phosphatase Secretion Mutants in Arabidopsis Thaliana
Jennifer Umpress 2001 Graduate Student Ph.D. in Plant Physiology Penn State Thesis: Phosphorus stress response in Arabidopsis
Gabriela Antunez De Mayolo 2003 Graduate Student Ph.D. in Integrated Life Sciences Penn State Thesis: Genetic engineering of Theobroma cacao and molecular studies on cacao defense responses; Past Position: Greifswald University (2010)
Melis Cakirer 2003 Graduate Student B.S. and M.S. in Food Sciences Penn State Thesis: Color as an indicator of flavanol content in the fresh seeds of Theobroma cacao L; Position: Colgate-Palmolive Company; 909 River Road, Piscataway, NJ 08855-1343 (2010), Email: melis_cakirer@colpal.com
J.D. Swanson 2005 Graduate Student Ph.D. in Integrated Biological Sciences Penn State Thesis: Flower development in Theobroma cacao L.: An assessment of morphological and molecular conservation of floral development between Arabidopsis thaliana and Theobroma cacao; Past Position: Professor of Biology and Class Dean, Department of Biology and Biomedical Sciences, Salve Regina University, 100 Ochre Point Ave, Newport RI 02840,Email: jswanson@uca.edu
J.P. Marelli 2008 Graduate Student Ph.D. in Integrated Biological Sciences Penn State Thesis: Solanum lycopersicum as a model system to study pathogenicity mechanisms of Moniliophthora perniciosa, the causal agent of witches' broom disease of Theobroma cacao; Past Position: Cocoa Physiologist; Mars Center For Cocoa Science; T: 55 73 3214-7906; M: 55 9116-3587; Fax: 55 73 3214-7905 (2010); Email: jean-philippe.marelli@effem.com
Carter Miller 2009 Graduate Student Ph.D. in Integrated Biological Sciences Penn State Thesis: An Integrated In Vitro and Greenhouse Orthotropic Clonal Propagation System for Theobroma Cacao L., Email: carter.r.miller@uol.com.br (2010)
Huan Xia 2009 Graduate Student Ph.D. in Integrated Biological Sciences Penn State Thesis: Structure and Function of Endosperm Starch from Maize Mutants Deficient in One or More Starch-branching Enzyme Isoform Activities; Position: Product Process Scientist R&D; MARS PETCARE US;T: 615-807-4234; Fax: 615-458-2743 (2010), Huan.Xia@effem.com
Zi Shi 2010 Graduate Student Ph.D. in Plant Biology Penn State Thesis: Functional analysis of Non Expressor of pr1 (NPR1) and its paralog NPR3 in Theobroma cacao and Arabidopsis thaliana. Past position: Scientist, Beijing Academy of Agricultural Forestry Science, shizi_520@msn.com
Rachel Melnick 2010 Graduate Student Ph.D. in Plant Pathology Penn State Thesis: Endophytic Bacillus SPP. of Theobroma Cacao: Ecology and Potential for Biological Control of Cacao Diseases
Yi Liu 2010 Graduate Student Ph.D. in Integrated Biological Sciences Penn State Thesis: Molecular Analysis of Genes Involved in the Synthesis of Proanthocyanidins in Theobroma in Cacao
Yufan Zhang 2014 Graduate Student Ph.D. in Plant Biology Penn State Thesis: Functional genomics of Theobroma cacao fatty acid biosynthesis: Convergence of fatty acid desaturation, embryo development, and defense signaling responses. Foreverkobe.zyf@gmail.com; formerly was a post doc at Princeton Uni.
Sergio Florez 2015 Graduate Student Ph.D. in Chemical Engineering Penn State Thesis: Improving Plant Propogation by Manipulating the Genetic and Physical Environment in Tissue Culture
Fister Drew 2016 Graduate Student Ph.D. in Genetics Penn State Thesis: Functional Genomics of the Theobroma Cacao Defense Response
Johann Restrepo Rubio 2020 Graduate Student M.S. Plant Sciences Penn State Thesis: Characterization of the Function of Selected Phytophthora Megakarya Effector Proteins During Infection of Theobroma Cacao
Knollenberg Benjamin 2020 Graduate Student Ph.D. Plant Biology Penn State Thesis: Characterization of Specialized Metabolites Involved in Black Pod Rot Resistance in Theobroma Cacao
Lina Tami- Barrera 2021 Graduate Student M.S. Dual title in Rural Sociology and International Agriculture & Development Penn State Thesis: Socioeconomic and Technological Factors Influencing Technology Adoption in Cacao Farms of Two Post-conflict Regions in Colombia
Winters Noah 2022 Graduate Student Ph.D. Ecology Penn State Thesis: Evolutionary and Functional Genetics of Disease Resistance in Tehobroma cacao and its Wild Relatives
Alejandro Gil Aguirre 2022 Graduate Student Ph.D. Education, Development, and Community Engagement Penn State Thesis: Technology adoption by cacao farmers in San Vicente de Chucuri, Santander: the role of Cadmium regulations
Jhony Benavides 2023 Graduate Student Ph.D. dual-title in Soil Science and International Agriculture and Development Penn State Thesis: Unmanned Aerial Vehicle Photogrammetry for Mapping Hydrologically Sensitive and Phosphorus Critical Source Areas
Zach Dashner 2024 Graduate Student Ph.D. Horticulture Penn State Thesis: Novel Biotechnological Approaches to Understand And Reduce Cadmium in the Iron Uptake Pathway of Theobroma Cacao
Daniel Guarín 2024 Graduate Student Ph.D. dual-title in Soil Science and International Agriculture and Development Penn State Thesis: The Interplay of Bedrock Geomorphology and Human Impact on Soil Health: A Multiscale Assessment
Jishnu Bhatt 2025 Graduate Student Ph.D. of Philosophy Penn State Thesis: Enhancing the Efficiency of Somatic Embryogenesis and Genetic Transformation in Theobroma cacao L.
Francisco Miguel Menéndez Burns 2025 Graduate Student Ph.D. Plant Biology Penn State Thesis: Genomics and Physiological Response to Cadmium and the Evolution of Heavy Metal ATPases in Theobroma cacao L.
David Hulce 2000 Staff Research Assistant
Lisa Renshaw-Gegg 2001 Staff Research Assistant, Lab Manager
Nur Sabrina Wahid 2010 Staff Research Assistant
Sharon Pishak 1996-2013 Staff Research Assistant, Lab Manager
Linda Miller 1999-2000 Staff Research Assistant, Lab Manager
Ann Young 1999-2010 Staff Research Assistant, Lab Manager
Nianying Wang 2002-2003 Staff Research Assistant
Melanie Schupp 2004-2005 Staff Research Assistant
Marissa Kenzakoski 2010-2013 Staff Research Assistant
Adam Clemens 2010-2013 Staff Research Assistant
Matthew Dommel 2012-2013 Staff Research Assistant
Teresa Soler 2014-2015 Staff Greenhouse/ lab Assistant
Andrea Murphy-Faust 2017-2018 Staff Research Assistant

Borlaug Fellow
First name Last Name Year(s) Home Country (from) Research
Monica Santos-Mendoza 2011 Mexico Transcriptomic and Motobolomic of Cocoa embriogenesis
Leidy Paola Delgadillo Duran 2020 Colombia Cd/ CAS9
Peter Aikpokpodion 2006-2007 Nigeria Genetic Diversity in Cacao (Theobroma cacao L.) Collections in Nigeria Determined by Phenotypic and Simple Sequence Repeat (SSR's) Markers

Fulbright Scholar
First name Last Name Year(s) Home Country (from) Research
Chia-Lock Tan 2006-2007 Malasia Molecular and Biotechnological Techniques for the Improvement of Cocoa Against Pests
Antonio Figueira 2007-2008 Brazil Global Analysis of Genes Differentially Expressed in the Theobroma Cacao Genotypes in Response to Witches Broom Infection
Nicolas Dr. Niemenak 2010-2011 Cameroon Maturation and germination physiology of cacao

Visiting Scientist
First name Last Name Year(s) Home Country (from) Research
Catalina Chapparo 2001 Colombia Cacao Somatic Embryogenesis for Colombian Varieties
Anna Muyiwa 2008 Nigeria Cacao, germplasm & germination
Yaleidis Mendez 2010 Puerto Rico Cacao somatic embryogenesis
Claudia Arevalo 2012 Venezuela Cacao Transformation
Dadzie Abu 2013 Ghana Disease resistance in cacao
Suzana Sato Guima 2015 Brazil Knowledge sharing
Lin Zhou 2015 China Defense gene in diversity population
Adrielle de Vasconcelos 2015 Brazil Knowledge sharing
Annelle Holder-John 2016 Trinidad & Tobago Cutting evaluation in two species of cacao
Jaime Osorio 2016 Colombia Cadium uptake in cacao
Gabriela Silva-Pumarada 2018 Puerto Rico Cacao
Fernando Santiago 2018 Puerto Rico Cacao
Thierry Legrave 2019 France CRISPR disease genes
Rosalia Velasquez-Salazar 2008-2009 Venezuela Cacao Somatic Embryogenesis/ Jatropha Somatic Embryogenesis
Luis Mejia-Franco 2009, 2012 Panama Cacao Endophytes
Desire Pokou 2013, 2016, 2017 Ivory Coast Disease resistance in cacao
Marieke Hilgers 2019-2020 Netherlands Functional genomics
Adriana Gallego 2106-2017 Colombia Polyphenol light effect

Graduate Students

Mark Guiltinan
1986
Graduate Student, Ph.D. Biology
University of California, Irvine
Thesis: The Isolation, Characterization and Intergeneric Transfer of Two Soybean (Glycine Max L.) Beta-tubulin Genes (TI Plasmid)


Charlene Chang
1993
Graduate Student, M.S. Plant Physiology
Penn State
Thesis: Optimization of in vitro regeneration of geranium (Pelargonium xhortorum L.H. Bailey) for development of a genetic transformation system.


Dane Fisher
1995
Graduate Student, Ph.D. in Genetics
Penn State
Thesis: Molecular genetic analysis of multiple isoforms of starch branching enzyme with emphasis on Zea mays L.; 


Xiping Niu
1995
Graduate Student, Ph.D. in Plant Physiology
Penn State
Thesis: DNA binding specificity and interactions with nucleosomal DNA of the plant bZip protein EmBP-1; 


Sashi Prabhakar
1995
Graduate Student, M.S. Plant Physiology
Penn State
Thesis: cDNA cloning and characterization of a EmBP-1 homologue in maize (Zea mays): A basic leucine zipper transcription factor; 


Ming Gao
1996
Graduate Student, Ph.D. in Plant Physiology
Penn State
Thesis: Molecular characterization of starch branching enzyme genes, SbeI, Sbe2b and Sbe2a in maize (Zea mays L.);


Siela Maximova
1996
Graduate Student, Ph.D. in Plant Physiology
Penn State
Thesis: Development of a genetic transformation system for apple; 


Kyung-Nam Kim
1997
Graduate Student, Ph.D. in Plant Physiology
Penn State
Thesis: Isolation and promoter analysis of the genes for starch branching enzyme in maize 


Abdoulaye Traore
1999
Graduate Student, Ph.D. in Food Science
Penn State
Thesis: Tissue culture and Transformation of cacao; 


Jennifer Tomscha
2001
Graduate Student, Ph.D. Horticulture
Penn State
Thesis: Phosphatase Secretion Mutants in Arabidopsis Thaliana


Jennifer Umpress
2001
Graduate Student, Ph.D. in Plant Physiology
Penn State
Thesis: Phosphorus stress response in Arabidopsis


Gabriela Antunez De Mayolo
2003
Graduate Student, Ph.D. in Integrated Life Sciences
Penn State
Thesis: Genetic engineering of Theobroma cacao and molecular studies on cacao defense responses; 


Melis Cakirer
2003
Graduate Student, B.S. and M.S. in Food Sciences
Penn State
Thesis: Color as an indicator of flavanol content in the fresh seeds of Theobroma cacao L; 


J.D. Swanson
2005
Graduate Student, Ph.D. in Integrated Biological Sciences
Penn State
Thesis: Flower development in Theobroma cacao L.: An assessment of morphological and molecular conservation of floral development between Arabidopsis thaliana and Theobroma cacao; 


J.P. Marelli
2008
Graduate Student, Ph.D. in Integrated Biological Sciences
Penn State
Thesis: Solanum lycopersicum as a model system to study pathogenicity mechanisms of Moniliophthora perniciosa, the causal agent of witches' broom disease of Theobroma cacao; 


Carter Miller
2009
Graduate Student, Ph.D. in Integrated Biological Sciences
Penn State
Thesis: An Integrated In Vitro and Greenhouse Orthotropic Clonal Propagation System for Theobroma Cacao L.,

Huan Xia
2009
Graduate Student, Ph.D. in Integrated Biological Sciences
Penn State
Thesis: Structure and Function of Endosperm Starch from Maize Mutants Deficient in One or More Starch-branching Enzyme Isoform Activities;


Zi Shi
2010
Graduate Student, Ph.D. in Plant Biology
Penn State
Thesis: Functional analysis of Non Expressor of pr1 (NPR1) and its paralog NPR3 in Theobroma cacao and Arabidopsis thaliana. 


Rachel Melnick
2010
Graduate Student, Ph.D. in Plant Pathology
Penn State
Thesis: Endophytic <i>Bacillus</i> SPP. of <i>Theobroma Cacao</i>: Ecology and Potential for Biological Control of Cacao Diseases


Yi Liu
2010
Graduate Student, Ph.D. in Integrated Biological Sciences
Penn State
Thesis: Molecular Analysis of Genes Involved in the Synthesis of Proanthocyanidins in Theobroma in Cacao


Yufan Zhang
2014
Graduate Student, Ph.D. in Plant Biology
Penn State
Thesis: Functional genomics of Theobroma cacao fatty acid biosynthesis: Convergence of fatty acid desaturation, embryo development, and defense signaling responses.


Sergio Florez
2015
Graduate Student, Ph.D. in Chemical Engineering
Penn State
Thesis: Improving Plant Propogation by Manipulating the Genetic and Physical Environment in Tissue Culture


Fister Drew
2016
Graduate Student, Ph.D. in Genetics
Penn State
Thesis: Functional Genomics of the Theobroma Cacao Defense Response


Johann Restrepo Rubio
2020
Graduate Student, M.S. Plant Sciences
Penn State
Thesis: Characterization of the Function of Selected Phytophthora Megakarya Effector Proteins During Infection of Theobroma Cacao


Knollenberg Benjamin
2020
Graduate Student, Ph.D. Plant Biology
Penn State
Thesis: Characterization of Specialized Metabolites Involved in Black Pod Rot Resistance in Theobroma Cacao


Lina Tami- Barrera
2021
Graduate Student, M.S. Dual title in Rural Sociology and International Agriculture &amp; Development
Penn State
Thesis: Socioeconomic and Technological Factors Influencing Technology Adoption in Cacao Farms of Two Post-conflict Regions in Colombia


Winters Noah
2022
Graduate Student, Ph.D. Ecology
Penn State
Thesis: Evolutionary and Functional Genetics of Disease Resistance in Tehobroma cacao and its Wild Relatives


Alejandro Gil Aguirre
2022
Graduate Student, Ph.D. Education, Development, and Community Engagement
Penn State
Thesis: Technology adoption by cacao farmers in San Vicente de Chucuri, Santander: the role of Cadmium regulations


Jhony Benavides
2023
Graduate Student, Ph.D. dual-title in Soil Science and International Agriculture and Development
Penn State
Thesis: Unmanned Aerial Vehicle Photogrammetry for Mapping Hydrologically Sensitive and Phosphorus Critical Source Areas


Zach Dashner
2024
Graduate Student, Ph.D. Horticulture
Penn State
Thesis: Novel Biotechnological Approaches to Understand And Reduce Cadmium in the Iron Uptake Pathway of Theobroma Cacao


Daniel Guarín
2024
Graduate Student, Ph.D. dual-title in Soil Science and International Agriculture and Development
Penn State
Thesis: The Interplay of Bedrock Geomorphology and Human Impact on Soil Health: A Multiscale Assessment


Jishnu Bhatt
2025
Graduate Student, Ph.D. of Philosophy
Penn State
Thesis: Enhancing the Efficiency of Somatic Embryogenesis and Genetic Transformation in Theobroma cacao L.


Francisco Miguel Menéndez Burns
2025
Graduate Student, Ph.D. Plant Biology
Penn State
Thesis: Genomics and Physiological Response to Cadmium and the Evolution of Heavy Metal ATPases in Theobroma cacao L.