Progress 10/01/08 to 09/30/09
Outputs
OUTPUTS: 2009 was a year of transitions for my laboratory. Each of the students and postdocs working on the Oenothera project (prior NSF grant) graduated or left for new positions, leaving the project to new postdoc Dr. Martin von Arx, who joined the lab from Neuchatel, Switzerland. Martin's first contribution to our program was to bring a new study organism, the hawkmoth Hyles lineata, into laboratory culture. This moth is the primary pollinator and herbivore on Oenothera plants, and will allow us to address a new suite of behavioral and ecological questions. Another transition was to the first full field season for the newly NSF-funded "red flower" project, which coordinates research teams in FL, NC and Ithaca focused on understanding the evolution of floral mimicry. This effort was led by postdoc Dr. Anne Gaskett, who collected critical data in FL and helped all teams standardize their experimental design, data collection and analysis. Finally, my most senior doctoral student (Joaquin Goyret) and postdoc (Dr. Rainee Kaczorowski) finished their work on foraging behavior by Manduca sexta and took new postdoctoral positions elsewhere. Their activities led to the funding of a new NSF grant on the behavioral responses of Manduca to relative humidity, which will support Dr. von Arx for part of 2010 and will allow Dr. Goyret to return to Cornell, if he chooses. Our activities led to the publication of 8 papers (see below), 7 meeting abstracts, numerous speaker invitations and public outreach activities. Three examples stand out. First, Dr. Kaczorowski presented two papers at the annual meeting of the Ecological Society of America, which is widely attended by pollination biologists. Rainee presented compelling data for the importance of floral volatiles in two different pollination systems (subtropical moth pollinated tobaccos and bumblebee pollinated alpine wildflowers). Second, doctoral student Holly Summers presented her unique data on population-wide scent sampling at the 4th Gordon Research Conference on plant volatiles, drawing attention to unprecedented variation in signal composition among neighboring plants. Finally, I presented an invited talk at a symposium on mutualism held at Harvard University, at which he challenged the current dogma on community-level plant-pollinator interactions by presenting chemical data that clearly structure such interactions. Critical proof-of-concept data were collected from field populations of Yucca plants (in TX) and fly-dispersed mosses (in Newfoundland) that will be used to prepare the next round of NSF and National Geographic funding proposals for 2010. PARTICIPANTS: Honors undergraduates Alison Seliger and Jessica Walden graduated with honors and moved to Boston, Alison to attend Harvard Dental School and Jessica to work as a lab technician at Tufts University and prepare her applications for doctoral programs in biological science. Rong Ma designed an honors project and will graduate in May 2010. Graduate student Joaquin Goyret defended his doctoral thesis in May 2009, having published 6 papers from his thesis research, and moved to Lund University, Sweden for postdoctoral work. Holly Summers progressed in her thesis work and submitted an NSF DDIG proposal in November 2009, whereas new doctoral student Paul Shamble joined the lab (co-mentored with Prof. Ron Hoy) in Sept. 2009. Postdoctoral fellow Dr. Rainee Kaczorowski left the lab in Sept. 2009 to take a postdoctoral position at the University of Arizona. Dr. Anne Gaskett completed her first year of postdoctoral work on our NSF-funded red flower project, then left Cornell to take a tenure track assistant professor position at Aukland University, New Zealand. I have hired Logan Jensen, a recent graduate from Warren Wilson College, NC to fill her position on the grant, but as a research assistant. In April 2009 we welcomed Dr. Martin von Arx as a new postdoctoral fellow in the lab, with one year of funding from the Swiss NSF. TARGET AUDIENCES: Not relevant to this project. PROJECT MODIFICATIONS: Not relevant to this project.
Impacts
The major impact of my research on my field this year took the form of a gathering of critical mass. I have steadily built the case for the widespread importance of floral chemistry, volatiles in particular, in generalized pollination systems more commonly studied by my colleagues. In 2009 I presented that case with the publication of an Annual Review of Ecology, Evolution and Systematics paper (Vol. 39: 549-569, Dec. 2008) and by guest editing a Special Feature in Functional Ecology (Oct. 2009, Vol. 23), including 7 invited papers on flower-pollinator interactions. I have seen an immediate impact from these activities, from high numbers of invitations as a seminar speaker to requests to train colleagues in my lab. I served as an external judge for a thesis at the Univ. of Toronto (July 2009), a professor-in-residence at the Univ. de Chile, Santiago (Oct. 2009) and an external reviewer for the Swedish Agricultural University in Alnarp, Sweden (Nov. 2009). In Aug. 2009 I chaired a symposium at the 4th Gordon Research Conference on plant volatiles in Oxford, UK, where these themes were further elucidated through the presentations of diverse participants. Plants' use of volatile compounds to manipulate their mutualists and antagonists is an idea that is beginning to take hold. My research explored several model systems in 2009, from the newly NSF-funded project investigating early spring-blooming, red flowers (e.g. Trilliums and pawpaws), to older projects on nectar-based pollination of Polemonium viscosum, a Rocky Mt. wildflower, to obligate mutualism between Yucca treculeana and its moth pollinators in southern TX. This work yielded exciting results, from the potential for microbial involvement in floral fermentation (in pawpaws) to the repellent aspects of strong scent produced by Polemonium flowers, which has an indirect effect on pollinator (bumble bee) mediated selection on floral shape in that system. We have prepared a manuscript describing this result for submission to a high impact journal. We also observed scale-dependent functions for floral scent in Yucca-yucca moth interactions, with orientation to scent away from the plant, but scent-triggered search behavior once the moth has landed on the flower spike. A second line of research represents a logical extension of our prior work on Manduca behavioral responses to environmental cues by asking how moths use relative humidity to find appropriate habitats and regulate their osmotic balance. This NSF-funded project (start date 1 Jan 2010) opens new possibilities for climate based research (with Dept. of Atmospheric Sciences at Cornell) and outreach activities from the Insectapalooza Entomology Open House to demonstrations at the Arizona-Sonora Desert Museum. One common problem among scientists is our specialization: meteorologists tend not to have training in life sciences, and few biologists truly understand how to interpret climatic data. Our new NSF funded project on relative humidity includes a plan to cross-train atmospheric science students in hawkmoth behavior and pollination ecology, including field work in the canyons of southeastern AZ.
Publications
- Goodrich, K.R., R.A. Raguso. 2009. The olfactory component of floral display in Asimina and Deeringothamnus (Annonaceae). New Phytologist 183: 457-469.
- Goyret, J., A. Kelber, M. Pfaff, R.A. Raguso. 2009. Flexible responses to visual and olfactory stimuli by foraging Manduca sexta: larval nutrition affects adult behaviour. Proceedings of the Royal Society (B) 276: 2739-2745.
- Raguso, R.A. 2009. Special feature: Floral scent in a whole-plant context: moving beyond pollinator attraction. Functional Ecology 23: 837-840
- Schlumpberger, B.O., A.A. Cocucci, M. More, A. Sercic, R.A. Raguso. 2009. Extreme variation in floral characters and its consequences for pollinator attraction among populations of an Andean cactus. Annals of Botany 103: 1489-1500.
- Jofre, J., F. Massardo, R. Rozzi, B. Goffinet, P. Marino, RA. Raguso, N. P. Navarro. 2009. Fenologia reproductiva del musgo Tayloria dubyi en las turberas de la Reserva de Biosfera Cabo de Hornos: un caso de interaccion entre musgo, dipterous y aves. Revista Chilena de Historia Natural: IN PRESS.
- Majetic, C.J., R.A. Raguso, T-L. Ashman. 2009. The sweet smell of success: floral scent affects pollinator attraction and seed fitness in Hesperis matronalis. Functional Ecology 23: 480-487.
- Marino, P., R.A. Raguso, B. Goffinet. 2009. The ecology and evolution of fly-dispersed dung mosses (Splachnaceae): manipulating insect behaviour through odour and visual cues. Symbiosis 47: 61-76.
- Wiemer, A., M. More, S. Benitez-Vieyra, A. Cocucci, R.A. Raguso, A. Sersic. 2009. A simple floral fragrance and unusual osmophore structure in Cyclopogon elatus (Orchidaceae). Plant Biology 11: 506-514.
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Progress 10/01/07 to 09/30/08
Outputs
OUTPUTS: My laboratory has aggressively pursued three projects this year. The first represented the final year of a 4 year NSF-funded project on floral scent variation in Oenothera plants, requiring two field teams to work from April-July in UT and WY. That grant ended as of 1Aug08 and we are preparing 4 manuscripts and a renewal application for continued funding on that project. Postdocs Derek Artz and Cristian Villagra completed their tenure in my lab during summer 08 and are now employed elsewhere as research associates. The second project involved experimental studies of Manduca moth behavior in the laboratory, with both artificial and natural flowers, concluding a previous NSF grant and laying the groundwork for a proposal submitted in July08 and under review at present. This work constitutes the PhD thesis of Joaquin Goyret and postdoctoral research of Rainee Kaczorowski. The third project began with my new NSF grant on 1July08, to study odor-color interactions in Trillium and similar plants. New postdoc Anne Gaskett will take the lead on this project over the next 3 years. During summer08, I co-chaired a symposium at the Int. Congress of Entomology in Durban, South Africa, helped organize the Int. Soc. of Behavioral Ecology congress on campus in Ithaca and gave an invited symposium talk at the Int. Soc. of Chemical Ecology meeting at Penn State. In October I gave a guest lecture in the Sensory Ecology short course at Lund University, Sweden. These experiences have led to several student recruiting opportunities, which will ensure the continued flow of new ideas through my laboratory. Beyond my own travels, my students have represented our lab and their own work at several international meetings this year (10 abstracts total), including Goyret's poster at the Invertebrate Vision Conference in Sweden, and posters by Kaczorowski and Artz at a symposium on pollination ecology held during the Ecological Soc. of America meeting in Milwaukee. Our work led to ten publications during 2008, including such high impact journals as Science (invited perspective), PNAS (invited paper), Annual Review of Ecology, Evolution and Systematics (invited review), and an invitation to edit a special feature on plant volatile ecology in the journal Functional Ecology. Clearly my lab's focus on plant volatile communication is beginning to reach a broader scientific audience. PARTICIPANTS: The final year of our NSF funded project on Oenothera featured projects by postdocs Derek Artz and Cristian Villagra, doctoral student Holly Summers, Honors undergraduate Jessica Walden and outgoing lab manager, Annie Simonin. Studies of Manduca sexta behavior continued with the doctoral thesis work of Joaquin Goyret and postdoc Rainee Kaczorowski, along with Honors undergraduate Alison Seliger. The new NSF-funded grant on red flowers began in July with the hire of postdoctoral fellow Anne Gaskett from Australia, who will remain with the lab through 2011. TARGET AUDIENCES: Not relevant to this project. PROJECT MODIFICATIONS: Not relevant to this project.
Impacts
Our research projects have been converging upon a similar message for several years: animals show flexible responses to sensory information, depending upon the other kinds of information available and their own prior experience and physiological condition. This message sounds self evident, but it has been generally ignored in behavioral research and is relevant to a broad spectrum of human interests, from integrated pest management in agriculture to the control of disease vectoring animals in public health, to the study of biological diversity and the maintenance of complexity in natural ecosystems. Another exciting and very basic outcome of our research is the idea that plants can manipulate animal behavior through the modification of floral odors and color signals that "tap into" the animals' basic biology. I promoted this idea in 2004 with an invited paper on the presence of odors in floral nectar, accompanied by a series of predictions about how plants might use such a feature strategically. This idea, tantamount to "plant behavior", has been emphasized in a recent Science paper by Ian Baldwin's group about nicotine in the nectar of wild tobaccos, for which I was invited to provide a perspective. Plant behavior will serve as the basis for a symposium which I will chair at the next Gordon Research Conference on Plant Volatiles (Aug. 2009, Oxford UK). Another impact from this research area has been the growing recognition from microbiologists that floral nectar often contains yeasts, that these microbes compete with pollinators for nectar sugar and that plants can either defend their flowers against yeasts or exploit them to optimize their reproductive needs (i.e. by making pollinators move further after a floral visit). Carlos Herrera's recent study of changes in floral nectar sugar by yeasts is a logical follow up to this idea, and has made an immediate impact on the pollination community. The impacts of my research on the professional development of my students have been tangible. Postdocs Derek Artz and Cristian Villagra left my lab after 3 years of experience working on the Oenothera project, including organizing field expeditions and data collection, mentoring younger students and developing their own ideas for grant and job applications. Both of them found employment as research associates working on similar themes. Anne Simonin, who joined me at Cornell after 3 years in my lab at USC, benefited greatly from her time as a lab manager here and has moved on to medical school in Charleston SC.
Publications
- Raguso, R.A. 2008. Wake up and smell the roses: the ecology and evolution of floral scent. Annual Review of Ecology, Evolution and Systematics 39: 549-569.
- Raguso, R.A. 2008. Start making scents: the challenge of integrating chemistry into pollination ecology. Entomologia Experimentalis et Applicata 128: 196-207.
- Schlumpberger, B.O., R.A. Raguso. 2008. Geographic variation in floral scent of Echinopsis ancistrophora (Cactaceae); evidence for constraints on hawkmoth attraction. Oikos 117: 801-814.
- Goyret, J., M. Pfaff, R.A. Raguso, A. Kelber. 2008. Why do Manduca sexta feed from white flowers Innate and learnt colour preferences in a hawkmoth. Naturwissenschaften: 95: 569-576.
- Goyret, J., P.M. Markwell, R.A. Raguso. 2008. Scale- and context-dependent effects of floral CO2 on nectar foraging by Manduca sexta. PNAS 105: 4565-4570.
- Wiemer, A., M. More, S. Benitez-Vieyra, A. Cocucci, R.A. Raguso, A. Sersic. 2008. A simple floral fragrance and unusual osmophore structure in Cyclopogon elatus(Orchidaceae). Plant Biology: doi:10.1111/j.1438-8677.2008.00140.x.
- Majetic, C.J., R.A. Raguso, T.-L. Ashman. 2008. The impact of biochemistry vs. population membership on floral scent profiles in colour polymorphic Hesperismatronalis. Annals of Botany: doi:10.1093/aob/mcn181.
- Raguso, R.A. 2008. Perspective: The 'invisible hand' of floral chemistry. Science 321: 1163-1164.
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Progress 10/01/06 to 09/30/07
Outputs
OUTPUTS: My CRIS project officially began on July 1, 2007, when I joined the NBB faculty at Cornell. Much of the remainder of 2007 was spent building my laboratory, identifying resources on the Ithaca campus and beginning to culture experimental organisms (Manduca moths, plant species in the genera Oenothera and Nicotiana). Pursuant to establishing an extramurally funded lab at Cornell, I submitted a proposal to NSF, in conjunction with co-PIs at Warren Wilson College (Asheville NC) and Georgia Southern University (Statesboro GA) in July 2007. The project was funded on Dec. 10, 2007, for $500,000, of which $350,000 will be awarded to Cornell, starting on 1 July 2008. The proposal outlined 5 alternative hypotheses to test regarding the evolution of red vs. white flowering Trillium plants in the forests of southeastern North America. I will extend the project to two additional Trillium species native to NY, which I will study locally with Cornell students. In October I chaired a
symposium on pollination and floral scent at the third Gordon Research Conference on Plant Volatiles, a conference that I co-founded in 1999. This meeting occurred in Switzerland and was attended by the world's most active researchers in this field. My symposium surveyed the progress made in the past decade in the integration of floral scent into pollination biology. The conference allowed me to recruit a postdoctoral scientist, Anne Gaskett, from MacQuarie University, Australia, to work on the Trillium project (2008-2011). Broader impacts of my research program took the form of outreach activities with a charter school near our field site in Moab, Grand Co. UT in June, a dinner with incoming Cornell freshmen in September, interviews with National Geographic and Science to provide commentary on Irene Terry's study of cycad pollination in Australia, and a subsequent one-week training workshop for Dr. Terry here at Cornell, in December. The Moab outreach programs, in conjunction with my
postdoc Dr. Derek Artz, presented the basics of pollination ecology to K-5 and high school students, including the plan to plant gardens of our study organism (Oenothera cespitosa) at these schools so that children could monitor pollinator and herbivore activity in suburban Moab and, potentially, develop web-based systems to communicate their findings. This is the kind of broader impact advocated by NSF, and is a logical outcome of our work with several volunteer gardeners and teachers from Moab. I am frequently asked to provide commentary on high impact studies in odor-mediated chemical ecology, which helps to publicize my field and recruit students to my laboratory. I have trained over 40 students and colleagues in the analysis of plant volatiles using gas chromatography-mass spectrometry since 1997, many of whom become co-authors in years to come. The study of plant volatile chemical ecology is growing rapidly, and my activities help to standardize methods and foster an environment
in which chemical approaches become accepted as mainstream tools in modern ecology.
PARTICIPANTS: Polemonium project (July, Colorado): Dr. Candace Galen, Univ. of Missouri, Dr. Rainee Kaczorowski, now postdoc in my lab. Floral CO2 project: Joaquin Goyret, PhD student in my lab. Oenothera floral variation project: Drs. Derek Artz and Cristian Villagra, postdocs, Annie Simonin, lab manager. Trillium project: collaborators, Dr. Amy Boyd (Warren Wilson College, Asheville NC), Dr. Michelle Zjhra, Georgia Southern University (Statesboro GA).
Impacts
My first research activity upon moving to Cornell was week-long collaborative project with Dr. Candace Galen (University of Missouri, Columbia) on the interplay between pollination and defense in the alpine wildflower Polemonium viscosum, in the tundra above Fairplay, Colorado. Dr. Galen pioneered the microevolutionary study of pollinator mediated selection on floral form and scent in these plants, but had never formally incorporated chemical analyses into her system. Polemonium floral odors are polar compounds that passively absorb into the nectar, so they have flavors as well as scents to visiting insects. Our experiment showed that 10-fold higher concentrations of scent were repellent to the flower's chief enemies, Formica ants, but had no impact on the guild of bees and flies that pollinate the plant. This exciting result suggests that floral characters may sometimes reflect defensive, as well as attractive qualities. Dr. Galen and my new postdoc, Rainee
Kaczorowski, are now poised to use chemical manipulations in further experiments with ant and bee behavior, as we develop these results into a grant proposal. My previous two NSF grants reached fruition this summer with the maturation of my students and our understanding of two new technologies. The first was to use a Licor CO2 analyzer to measure and manipulate CO2 emissions from newly opening, night blooming flowers visited by our study organism, the hawkmoth Manduca sexta. My doctoral student, Joaquin Goyret, demonstrated that Manduca moths utilize CO2 as an odor, tracking it to find plants from a distance, but ignore it once they arrive at the flower, using other cues to probe for nectar. Interestingly, females only respond to CO2 in the presence of a host plant odor, suggesting that feeding and egg laying are inextricably combined tasks for them. This is the first demonstration of the scale- and context-dependent roles for CO2 in pollination biology, and will be published this
spring in PNAS. The second was to use a portable fast-GC-surface acoustic wave detector device to survey hundreds of individual Oenothera cespitosa flowers in Moab UT and Jackson WY, for variation in floral scent chemistry. Population surveys of floral chemistry performed by my postdoc Derek Artz and lab manager Annie Simonin have not been undertaken before at this scale, and we learned that at least 5 different "odor morphs" are present in each habitat, with some of them being quite rare. The up coming field season will allow us to test whether these odors are associated with differences in herbivore damage, pollinator visitation or pathogen infection.
Publications
- Theis, N.B., M. Lerdau, M., Raguso, R.A. 2007. The challenge of attracting pollinators while evading floral herbivores: Patterns of fragrance emission in Cirsium arvense and Cirsium repandum (Asteraceae). Int. J. Plant Sci. 168: 587-601.
- Majetic, C.J., Raguso, R.A., Tonsor, S.J., Ashman, T.L. 2007. Flower color-flower scent associations in polymorphic Hesperis matronalis (Brassicaceae). Phytochem. 68: 865-874.
- Raguso, R.A., Kelber, A., Pfaff, M., Levin, R.A. and McDade, L.A. 2007. Floral biology of North American Oenothera sect. Lavauxia (Onagraceae): Advertisements, rewards and extreme variation in floral depth. Ann. MO Bot. Gard. 94: 236-257.
- Raguso, R.A., Ojeda-Avila, T., Desai, S., Jurkiewicz, M.K. and Woods, H.A. 2007. The influence of larval diet on adult feeding behaviour in the tobacco hornworm moth, Manduca sexta. J. Insect Phys. 53: 923-932.
- Goyret, J., Markwell, P.M, Raguso, R.A. 2007. The effect of olfactory and visual stimuli decoupling on the foraging behavior of Manduca sexta. J. Expt. Biol. 210: 1398-1405.
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