S. Feldman: Research Interests and Previous Projects
Interactions between species can profoundly affect whether populations persist, grow, or become extinct. These interactions can also determine which organisms live together in communities. Still, few studies have linked species interactions with effects on populations and communities. In my lab, I study effects of species interactions on individual fitness, population dynamics and communities. Through my research, I aim to elucidate the influence of human-induced changes, such as habitat fragmentation and introduced species, on the behavior and population dynamics of species involved in multi-species interactions. I address these issues with a combination of field and lab experiments, statistical and mathematical models, culturing techniques, and molecular tools. I study a variety of systems, including multiple plant species that share pollinators, plants with fungal pathogens and insect vectors, insect herbivores and their predators, plant-associated endophytic fungi, and fungal viruses. My research involving fungi and viruses contributes to the rapidly developing fields of microbial and virus ecology, which help us gain a broader understanding of species diversity and species interactions in these extremely diverse and understudied groups. Thus far, my work has addressed three main questions. (1) What factors enable plant-pollinator mutualisms to persist when plants are at low densities? (2) What are the behavioral, fitness, and population consequences of competitors on mutualisms involving plant pathogens and their vectors? (3) Do diverse communities of fungal endophytes support even more diverse communities of fungal viruses? To illustrate my approaches to research, I will describe my work in each of these areas below.
(1) What factors enable plant-pollinator mutualisms to persist when
plants are at low densities?
Mutualisms are relationships in which two species have higher fitness when interacting than they would alone. Mutualisms may be particularly susceptible to extinction in rapidly changing or highly disturbed environments. One mechanism that may allow pollination mutualisms to persist is pollination facilitation, a process by which reproductive success of a rare plant species increases in the presence of a second species due to increased visits from shared pollinators. Plants occurring at low densities potentially suffer reduced visitation or pollination success, so facilitation might be critical to population persistence. I explored this question for my dissertation work (with Dr. W. F. Morris at Duke University), using a variety of systems and approaches, including mathematical models and field experiments. Undergraduate field assistants helped conduct these field experiments.
From two separate field
experiments with two different focal plants, Brassica rapa and Piriqueta
caroliniana, I found strong positive effects of increasing plant density on
pollinator visitation (measured as visits to plants per hour and visits to
plants per foraging bout) and reproductive success (seed production). In
addition, evidence from a density-dependent projection matrix model for
population growth in P. caroliniana demonstrates that populations may
grow more slowly at low densities due to reduced seed production, but do not
decline (Allee effects occur but are weak).
To determine when
pollination facilitation might rescue plants from negative effects of low
density, I conducted experiments using Coreopsis leavenworthii, a
co-flowering species that shares pollinators with P. caroliniana. I
found that visitation, pollen receipt, and reproductive success of P. caroliniana
remain unaffected in the presence of the co-flowering species. Thus,
pollination facilitation may not occur when plants simply share pollinators and
co-occur at low densities—other criteria are important. Evidence from a
mathematical model I developed (with Dr. W. F. Morris and Dr. W. G. Wilson)
suggests that one plant species can also facilitate another’s pollination when
the number of pollinator visits to patches of plants per unit time (the
aggregative response) accelerates at low densities. To my knowledge, this
mechanism by which facilitation might occur has not been addressed previously.
However, in a second experiment with Brassica rapa, the pollinator
aggregative response did not accelerate at low densities. Although pollination
facilitation has been found in several systems, and occurs by several different
mechanisms (e.g. through disproportionate increases in visits to individual
plants or through maintenance of larger pollinator populations in more diverse
plant communities), facilitation may be unlikely to occur through increases in
visits to patches of plants.
At the University of Wisconsin – Stevens Point, I have been addressing a related question in collaboration with (and funded by) the Wisconsin Department of Natural Resources, working with undergraduate students to collect data with which I can develop a demographic model of a rare wetland plant (Oxytropis campestris var. chartacea, or Fassett’s Locoweed). This plant occurs at extremely varied densities on lake shorelines where water levels fluctuate dramatically over decades. Plants exposed for long periods of time may face competition from other species, and plants inundated by water likely die, leaving only a seed bank to recolonize when water levels recede. Thus, I have incorporated into the model both demographic data from plants at high and low densities, as well as data from experiments to characterize the seed bank, germination rates, and seedling establishment. Very few demographic models have included effects of seed banks on population persistence.
experiments with mathematical models enabled me to use field data to predict
population dynamics under different scenarios. In general, this
interdisciplinary approach is necessary for understanding complex processes
that are difficult to measure directly. In the future, I hope to continue to
address population-level consequences of plant-pollinator interactions,
including other factors that may cause population decline, such as
self-pollination in outcrossing species.
(2) What are the
behavioral, fitness, and population consequences of competitors on mutualisms
involving plant pathogens and their vectors?
In almost all known mutualisms, additional species exploit or compete with one
or both partners, raising the possibility that exploiters could erode the
mutualism over ecological or evolutionary time. However, the consequences of
exploitation for most mutualisms are not well understood. I have been
developing a research project to address this question in the context of a
system of multi-species interactions involving Claviceps, a genus of
fungal pathogens that infect grasses. In the future, I plan to combine
fieldwork, laboratory/greenhouse experiments, and molecular work to study the
effects of a common pathogen (Claviceps paspali) on the fitness of two
invasive grass species (Paspalum notatum and P. dilatatum) that
are hosts to the pathogen. C. paspali can severely damage grain crops
and poison livestock that eat infected forage grasses. However, effects of this
pathogen on whole-plant fitness remain largely unknown. Claviceps
species are likely involved in mutualistic interactions with insect vectors.
During infections, C. paspali translocates sugars from its host grasses,
producing sugar-rich exudates (laden with spores) that may provide a major food
source to the hundreds of species of diverse insects in several orders that
visit infected grass inflorescences and disperse fungal spores. Some of the
insects attracted to infected inflorescences are capable of spreading fungal
spores, thereby potentially spreading the disease. Thus, the fungal pathogen Claviceps
paspali and its insect vectors are likely mutualists.
Other fungal species, including Fusarium heterosporum, may exploit the
mutualism between C. paspali and its insect vectors. F. heterosporum
grows on the exudates produced by C. paspali, thereby potentially
affecting fitness of the pathogen while also decreasing the potential reward
reaped by insects that visit inflorescences infected by both fungi. F.
heterosporum may also change the chemistry of the system, affecting insect
attraction to infected inflorescences or fitness of insects that feed on the
exudates. Further, several species of plant-associated fungi live inside the
leaves and stems of Paspalum species that are hosts of C. paspali,
with unknown effects on fitness of host grasses, C. paspali, F.
heterosporum, or insect vectors. I have submitted a paper coauthored by Dr.
H. E. O’Brien and Dr. A. E. Arnold demonstrating that moths carry spores of F.
heterosporum and fungal endophytes of Paspalum species. As a future
research project, I hope to use this system as a model for insect-vectored
pathogens of plants. Also, I believe that many questions in this system are
conducive to involving undergraduates in research.
(3) Do diverse communities of fungal endophytes support even more diverse communities of fungal viruses?
In many systems, parasitic and mutualistic symbionts are incredibly diverse. Although researchers have begun to characterize patterns of fungal diversity, little is known about the symbionts of fungi. Virtually all plant (and
perhaps animal) species harbor pathogenic or mutualistic fungi in their
tissues. Viruses of these fungi have the potential to affect parasitism by
changing virulence of fungal pathogens or mutualisms by altering host tolerance
to environmental stress. However, almost nothing is known about fungal viruses
in an ecological context. In addition, although the “shotgun” sequencing
approach has illuminated extremely diverse and previously unknown fungi,
bacteria and viruses, it is often difficult to determine the associations between
individual species from these presence/absence data. As part of my postdoctoral
research (part of a large NSF-EPSCoR funded project on plant virus
biodiversity, with Dr. M. J. Roossinck), I am conducting a study of fungal
virus prevalence and diversity in a tall grass prairie ecosystem in Oklahoma.
In this system, fungi in diverse genera are hosts to viruses in more than 5
families. All of the 19 viruses I encountered so far (from a sample of 200
plant-associated fungi) are likely to be novel, and some are not closely related to any known viruses. Fungal viruses are likely to be more diverse than their fungal hosts, because multiple virus types were found in each of the most common fungal taxa. Also, rescaled rarefaction curves suggest that with additonal sampling, we are likely to recover additional new virus taxa at higher rates than new fungal taxa.
hope to develop a system to study effects of some of these
fungal viruses on their fungal hosts (e.g., Alternaria alternata and Stemphylium
solani), and on plants associated with these fungi: the parasitic plant Cuscuta
cuspidata and one of its host plants, Ambrosia psilostachya. To conduct this work, I used several molecular tools, including DNA extraction, polymerase chain reaction (PCR), sterile culture techniques, double-stranded RNA extractions, reverse-transcription PCR to obtain mycovirus sequences, and phylogenetic tools (for classifying fungi and viruses). I hope to continue working to address effects of fungal viruses on their host fungi and on interactions between fungi and their associated plant hosts.
In my lab at the University of Wisconsin Stevens Point, I worked with undergraduate students to develop projects to address several other diverse questions, including predator responses to eyespots on caterpillars that may mimic snakes, and competitive hierarchies and colonization-competition tradeoffs in endophytic fungi co-occurring within tree branches.
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