Educational Path & Background
I have been fortunate to learn from inspiring, patient and excellent scientists. I met some by chance, others by following my interests, but all of them helped me to find my own voice as a biologist and introduced me to the community of scientists to which I now belong. Here I recognize their contributions to my education with gratitude. My first research experience came during two summers (1982-83) as a technician in Darcy Kelley's Xenopus frog lab at Columbia University. I was just a high school student, but Darcy introduced me to so many ideas, techniques and people (including John Hildebrand, see below) that would later become central to my life. She also introduced me to the concept of research centers like the Marine Biological Laboratory at Woods Hole, MA, where she taught each summer, a place described with great affection by Lewis Thomas in "The lives of a cell".
[Left, Kelley lab members at Woods Hole, MA, July 1983]
I have been fortunate to learn from inspiring, patient and excellent scientists. I met some by chance, others by following my interests, but all of them helped me to find my own voice as a biologist and introduced me to the community of scientists to which I now belong. Here I recognize their contributions to my education with gratitude. My first research experience came during two summers (1982-83) as a technician in Darcy Kelley's Xenopus frog lab at Columbia University. I was just a high school student, but Darcy introduced me to so many ideas, techniques and people (including John Hildebrand, see below) that would later become central to my life. She also introduced me to the concept of research centers like the Marine Biological Laboratory at Woods Hole, MA, where she taught each summer, a place described with great affection by Lewis Thomas in "The lives of a cell".
[Left, Kelley lab members at Woods Hole, MA, July 1983]
I attended Yale as an undergraduate (1983-87) to study butterflies with Charles Remington, a co-founder of the Lepidopterists' Society and an influential figure in the development of butterflies as model systems for the study of evolution and population genetics. As my first mentor, Charles introduced me to the culinary delights of New Haven, challenged me to expand my interests beyond butterflies (to moths, aquatic insects, biogeography, chemical ecology) and slyly motivated me to travel and gain field experiences of my own by invoking the accomplishments of my predecessors at Yale, including Lincoln Brower, Naomi Pierce, Bob Pyle, Larry Gall, Ward Watt and Francie Chew. I can only think of him when I see a bolo tie!
[Left, Charles Remington in Yale's Peabody Museum, courtesy of Science magazine, April 1992]
[Left, Charles Remington in Yale's Peabody Museum, courtesy of Science magazine, April 1992]
In summer 1985, I had two great research opportunities. The first came at Mountain Lake Biological Station near Blacksburg, VA, where I worked with Beverly Rathcke and Leslie Real. Their NSF-funded study of nectar variance in heath balds (mountain top communities of Kalmia and Vaccinium shrubs) focused on risk aversion by bee pollinators. The hard physical labor of data collection, debates over experimental design, intense social experiences and beautiful setting of Mt. Lake were stimulating, and the weeks passed quickly. I realized how little I knew about plants and became motivated to learn more about pollination.
[Right, Beverly Rathcke, Bald Knob, Mt. Lake Biological Station, Pembroke, VA, June 1985]
[Right, Beverly Rathcke, Bald Knob, Mt. Lake Biological Station, Pembroke, VA, June 1985]
Leaving the southern Appalachians, I flew to Veracruz, Mexico to join Carol Horvitz and Doug Schemske on another NSF-funded project, this one focused on plant-pollinator-ant ecological networks in the tropical understory herb, Callathea ovandensis. Each day we drove past the Olmec head in San Andres Tuxtla to our rainforest field site in the crater of a small volcano at Laguna Encantada, and returned each evening to the delightful town of Catemaco, with its salsa music, cowboy cuisine and brujo folklore. I was overwhelmed by tropical biodiversity and my first immersion in the Spanish language, and initiated a butterfly survey that would become my first publication.
[Left: 20-year old REU student scoring pollinators after an army ant raid, July 1985]
[Left: 20-year old REU student scoring pollinators after an army ant raid, July 1985]
I returned to Yale motivated to become a field biologist and to clean up my act academically. In retrospect I am astonished to have had the chance to learn from four terrific ecologists in one summer, each inspiring in their own ways. I was also very fortunate to have worked on such conceptually ground-breaking projects. Pollinator risk-sensitivity remains an important question decades later, as does the use of path analysis-like modeling approaches to untangle complex plant-pollinator interactions in a whole-plant context. Although I didn't realize it then, the summer of '85 set me on my current path.
My eventual honors research project, initiated with Rick Harrison and completed in Jeff Powell's lab, was a test of geographic isolation and genetic drift using allozyme and morphometric data from Colias interior butterfly populations separated by 8-12,000 generations due to the retreat of the Laurentides Ice Sheet in eastern North America. I won my first small grant, traveled from Maine to West Virginia to collect pink-edged sulphur butterflies and burned through a LOT of starch gel electrophoresis. But in the end, I failed to reject my null hypothesis and experienced a deep frustration with population genetics, which led me to take time away from science and, eventually, to change fields.
[Right, Ward Watt injecting a Colias butterfly flight muscle sample for HPLC analysis, Stanford Biology Dept. Bulletin, 1988]
Nevertheless, this project introduced me to Ward Watt, whose group studied the evolutionary genetics and functional ecology of Colias butterflies at Stanford University. Ward hired me as his lab technician for 2 years (1988-89) at Stanford, where I learned HPLC and PAGE, cared for thousands of caterpillars and decided that I should go on to graduate school. Ward always made time to talk candidly about academic science as a career path. I doubt that I would be a professor today if not for his guidance and care, and my mechanistic bias reflects his commitment to integrating between different levels of analysis.
[Right, Ward Watt injecting a Colias butterfly flight muscle sample for HPLC analysis, Stanford Biology Dept. Bulletin, 1988]
Nevertheless, this project introduced me to Ward Watt, whose group studied the evolutionary genetics and functional ecology of Colias butterflies at Stanford University. Ward hired me as his lab technician for 2 years (1988-89) at Stanford, where I learned HPLC and PAGE, cared for thousands of caterpillars and decided that I should go on to graduate school. Ward always made time to talk candidly about academic science as a career path. I doubt that I would be a professor today if not for his guidance and care, and my mechanistic bias reflects his commitment to integrating between different levels of analysis.
A seminar by Ward's colleague Leslie Gottlieb fundamentally altered my ideas about science through his explicit use of the hypothetico-deductive method to frame his research questions and his choice of fast-growing annual Clarkia plants as a genetically tractable, ecologically interesting model system for studying plant evolution. I was determined to bring something new to the study of insect-plant coevolution, and reasoned that the next stage of my education should feature the plant point of view. In 1989, I moved to Ann Arbor to begin my doctoral studies with Eran Pichersky at the University of Michigan. Eran was a young assistant professor who had studied gene duplication in Clarkia with Leslie at UC Davis and was interested in the evolution
of novel plant traits. [Right, Leslie Gottlieb (L) and Eran Pichersky (R), April 1991, Ann Arbor MI]
of novel plant traits. [Right, Leslie Gottlieb (L) and Eran Pichersky (R), April 1991, Ann Arbor MI]
Eran suggested that I study the de novo evolution of floral scent associated with a shift from bee to hawkmoth pollination in Clarkia breweri for my thesis. At the time I was taking a Plant-Insect Interactions seminar with Bev Rathcke, whom I already knew from Mt. Lake, and Michael Martin, a chemical ecologist who studied the nutritional ecology of leafcutter ants and caterpillars. My assignment was to assess the importance of floral scent in pollination, but the literature was largely empty. Aside from classic studies of sexually deceptive orchids and euglossine bees, and Heidi Dobson's pioneering work on pollen volatiles, nearly nothing was known about the genetics, bio-chemistry, ecology or evolution of floral scent. Anything that I found would be NEW!
[Above, Eran Pichersky visits a Clarkia breweri population, Del Puerto Canyon, CA, May 1990]
[Above, Eran Pichersky visits a Clarkia breweri population, Del Puerto Canyon, CA, May 1990]
Now my task was to learn how to collect and analyze floral volatiles, which nobody at Michigan was doing. With advice from Heidi Dobson, Olle Pellmyr, Mark Whitten and Roman Kaiser, I slowly mastered GC-MS working midnight-to-dawn shifts on an ancient Finnigan quadrupole in the UM Chemistry Dept. My epiphany came when I realized that a small GC peak whose mass spectrum suggested 2-methoxy-4-(2-propenyl) phenol or "eugenol" as a structure was actually clove oil, which made flowers of C. breweri smell "spicy" to my nose! Learning to equate mass spectra with odors that pleased or disgusted me was like decoding the script of an ancient language, and made learning organic chemistry a lot more fun. Patterns began to emerge from our data, such as a duplicate gene whose product methylates eugenol or isoeugenol in certain populations of C. breweri, which Jihong Wang would then study for her thesis. The pace of discovery doubled when Natalia Dudareva brought her RNA skills to Eran's lab. Their work has altered the landscape of plant volatile metabolism, and I feel fortunate to have explored the first blind alleys with them. Although I never became a plant molecular biologist, Eran patiently trained me to be a skeptical scientist, to respect data over theory, to organize my mind and my life.
[Clockwise from left: Clarkia breweri, C. concinna and their F1 hybrid, tracking the genetic control of linalool]
[Clockwise from left: Clarkia breweri, C. concinna and their F1 hybrid, tracking the genetic control of linalool]
When I started giving talks about how flowers make volatiles, most of the questions I fielded were about how pollinators smell them! During visits to his lab at the USDA-ARS in Albany, CA, Douglas Light taught me how to measure "electroantennograms", the antennal responses of hawkmoths to floral odors. This seemed like an exciting direction for postdoctoral research, a way to return to moths with a new set of tools. Doug suggested that I visit the University of Arizona, where John Hildebrand headed a division of neuroscientists who used the tobacco hornworm moth, Manduca sexta, as a model system for studying sensory organization. I had met John 13 years earlier at Woods Hole, and knew him to be a gregarious and energetic champion of insect olfactory neurophysiology. His interest in expanding models of olfaction to include plant volatiles as well as sex pheromones made me feel welcome as my research shifted direction yet again.
[Left, Doug Light at his EAG rig, USDA-ARS Albany CA, May 1994]
[Left, Doug Light at his EAG rig, USDA-ARS Albany CA, May 1994]
[Above; Mark Willis (L), sunset at the Arizona-Sonora Desert Museum (C), John Hildebrand & Gail Burd (R)]
My postdoctoral work in Tucson was funded through the Center for Insect Science (CIS), and while based in the Hildebrand lab I had two additional mentors: Mark Willis, a Jedi master of wind-tunnel behavioral assays with moths, and Reg Chapman, an eminent insect physiologist and author of the classic text " The Insects; structure and function". Mark and I worked together after sunset at the Arizona-Sonora Desert Museum, one of the truly magical places in my life. Many of our experiments were hatched during these evenings as we filmed wild hawkmoths visiting Datura flowers in the pollinator gardens. Tutorials with Reg convinced me that their feeding behavior had to be multimodal. John Alcock's evocative use of hypothesis trees in his "Animal Behavior" textbook inspired the design of our experiments, in which Mark and I demonstrated synergism between floral scent and visual display in nectar feeding by Manduca sexta, by experimentally decoupling and recombining these stimuli. These were simple yet satisfying ethological experiments, which generated many new questions about the sensory world of Manduca sexta, some of which continue to be studied by members of John Hildebrand's group as well as my own students.
My postdoctoral work in Tucson was funded through the Center for Insect Science (CIS), and while based in the Hildebrand lab I had two additional mentors: Mark Willis, a Jedi master of wind-tunnel behavioral assays with moths, and Reg Chapman, an eminent insect physiologist and author of the classic text " The Insects; structure and function". Mark and I worked together after sunset at the Arizona-Sonora Desert Museum, one of the truly magical places in my life. Many of our experiments were hatched during these evenings as we filmed wild hawkmoths visiting Datura flowers in the pollinator gardens. Tutorials with Reg convinced me that their feeding behavior had to be multimodal. John Alcock's evocative use of hypothesis trees in his "Animal Behavior" textbook inspired the design of our experiments, in which Mark and I demonstrated synergism between floral scent and visual display in nectar feeding by Manduca sexta, by experimentally decoupling and recombining these stimuli. These were simple yet satisfying ethological experiments, which generated many new questions about the sensory world of Manduca sexta, some of which continue to be studied by members of John Hildebrand's group as well as my own students.
[Above, Lucinda McDade (L) at the California Botanic Garden, Judie Bronstein (C) and Goggy Davidowitz (R)]
Like many in my cohort, I did not want to leave Tucson, where beautiful landscapes, a thriving artistic community, authentic Mexican cuisine and a critical mass of insect scientists made my postdoctoral years delightful. Beyond the Division of Neurobiology, I enjoyed interactions with Judie Bronstein and Goggy Davidowitz, with whom I still collaborate. During this time (1996-99) Tucson was ground-zero for the phylogenetic revolution, and I gravitated to Lucinda McDade, whose keen insights, generous mentorship and wicked humor attracted a crowd to her lab and the UA herbarium. Lucinda, her student Rachel Levin, and I puzzled over how biosynthetically correlated floral volatiles could be mapped onto phylogenetic relationships. This question fueled my second postdoc through an NSF training grant in Biodiversification to the Dept. of Ecology and Evolution, and later generated my first NSF grant (with Lucinda) and Rachel's doctoral thesis, focusing on three plant lineages in which hawkmoth pollination had been gained and lost. This project provided the transition to my first job, at the University of South Carolina.
Like many in my cohort, I did not want to leave Tucson, where beautiful landscapes, a thriving artistic community, authentic Mexican cuisine and a critical mass of insect scientists made my postdoctoral years delightful. Beyond the Division of Neurobiology, I enjoyed interactions with Judie Bronstein and Goggy Davidowitz, with whom I still collaborate. During this time (1996-99) Tucson was ground-zero for the phylogenetic revolution, and I gravitated to Lucinda McDade, whose keen insights, generous mentorship and wicked humor attracted a crowd to her lab and the UA herbarium. Lucinda, her student Rachel Levin, and I puzzled over how biosynthetically correlated floral volatiles could be mapped onto phylogenetic relationships. This question fueled my second postdoc through an NSF training grant in Biodiversification to the Dept. of Ecology and Evolution, and later generated my first NSF grant (with Lucinda) and Rachel's doctoral thesis, focusing on three plant lineages in which hawkmoth pollination had been gained and lost. This project provided the transition to my first job, at the University of South Carolina.
I'll conclude with a vivid memory of a turning point in my professional life. In 1998 I was struggling to land a job, running out of funding, and felt that I had fallen into the cracks between several fields. Gunnar Bergstrom was visiting Tucson and I joined him, John Hildebrand and John Law, the CIS director, for dinner. I asked Gunnar why there had never been a meeting or symposium focused on floral scent, to which he responded that his group had studied floral volatiles (especially in sexually deceptive Ophrys orchids) to learn more about bees, rather than to understand flowers per se. John Law, who was a board member of the Gordon Research Organization, suggested that Gunnar and I apply to establish a new Gordon Research Conference on Floral Volatiles, which we did! In fact, John Hildebrand established a new GRC for Neuroethology during the same year, which we celebrated in back-to-back meetings at Queens' College, Oxford, UK in September 1999.
One of the most thrilling moments of my life was walking onstage to welcome the attendees of our first conference, including many of the legendary chemists whose papers I had read over the years, but I had never met. Gunnar, Heidi Dobson, Jette Knudsen and I spent a year inviting these speakers, raising funds and stitching together a program to help create a new field. The cultural clashes, instant collaborations and intellectual energy resulting from that conference were unforgettable, and set the course for the next stage of my life, as a young professor of chemical ecology. Here's to you, John Law!