Source: WASHINGTON STATE UNIVERSITY submitted to NRP
TRACKING WITHIN-VECTOR PHYTOPLASMA DYNAMICS TO OPTIMIZE CONTROL
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
1026513
Grant No.
2021-67034-35173
Cumulative Award Amt.
$164,960.00
Proposal No.
2020-10532
Multistate No.
(N/A)
Project Start Date
Jun 15, 2021
Project End Date
Jun 14, 2023
Grant Year
2021
Program Code
[A7201]- AFRI Post Doctoral Fellowships
Recipient Organization
WASHINGTON STATE UNIVERSITY
240 FRENCH ADMINISTRATION BLDG
PULLMAN,WA 99164-0001
Performing Department
Entomology
Non Technical Summary
X-disease phytoplasma (Candidatus Phytoplasma pruni), the causal organism of X-disease in stone fruits or "buckskin" and "little cherry disease" in cherries, recently re-emerged in the US and is currently devastating cherry production in the Pacific Northwest. Diseased trees never recover, therefore control methods rely on infected tree removal and broad-spectrum chemical control of the leafhopper vectors. Over 1,000 acres of trees have been removed in the last five years and $18 million lost in revenue in 2020 alone due to X-disease.Research on leafhopper-X-disease phytoplasma interactions is nearly a half-century old leaving imprecise epidemiological knowledge, and hindering outbreak mitigation efforts. One interesting finding is the phytoplasma reduces the leafhopper's lifespan; therefore, enhancing the phytoplasma's virulence through coinfection with another insect pathogencould reduce vector capacity. This project utilizes a combination of molecular, physiological, and ecological approaches to describe and understand the disease cycle of X-disease within the leafhopper vectors and develop sustainable vector control techniques to limit transmission and outbreaks.Through laboratory and greenhouse experiments I will describe the X-disease infection cycle within the leafhopper vector from acquisition to transmission in both X-disease phytoplasma only infected leafhoppers and leafhoppers coinfected with a fungus. Specifically, I will use probes to fluorescently mark pathogen DNA within the vector at various time points post acquisition to visualize and determine the pathway of infection, time to the vector becoming infectious, and interactions of two pathogens within a host. The results from these experiments will determine how long it takes for the vector leafhoppers to become infectious after uptake of the pathogen which is critical to informing X-disease management strategies. Understanding the pathway of infection provides insights into phytoplasma disease ecology and can open new avenues for control options within the leafhopper vector. Assessing effects of coinfection will elucidate the effects of microbial control agents on the vector and X-disease phytoplasma. Accomplishing these objectives will build the foundation for sustainable X-disease management programs and provide insight into other arthropod vectored phytoplasmas.
Animal Health Component
40%
Research Effort Categories
Basic
50%
Applied
40%
Developmental
10%
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
2111112113030%
2121112104040%
2151112107010%
2161112113020%
Goals / Objectives
Project, Training, and Career Development Goals and Objectives:Goal 1: Expand into new dsciplines and pest system to develop interdisciplinary skills working with an insect-vectored pathogen to gain experience in insect physiology, disease ecology, and plant pathology. Objective 1: Create histological cross-sections of Colladonus reductus leafhoppers under guidance of Dr. Harper. 2: Dissect and mount leafhopper alimentary canal, and primary and accessory salivary glands under guidance of Dr. Cooper. 3: Describe the X-disease infection cycle within the leafhopper vector through insect DNA probing the dissected tissues and cross-sections. 4: Describe effects of co-infection with fungal entomopathegen and X-disease phytoplasma through histological cross-sections and dissections. 5: Determine incubation period, and daily mortality of infected vs uninfected leafhoppers to determine pathogen transmission probability.Goal 2: Enhance skills in molecular techniques to support ecological research and address grower challenges. Objective 1: Collect leafhopper and cherry tree tissueand conduct PCR test for phytoplasma infection. 2: Create and use DNA or RNA fluorescent probes to hybridize and visually observe pathogens in tissue.Goal 3: Gain experience communicating, teaching, and mentoring. Objective 1: Publish two scientific manuscripts from the research proposed. 2: Lead a discussion in a University Seminar on Career Development. 3: Assist in mentoring Dr. Northfield's graduate students, and lab technicians.MilestonesDevelop methods to use molecular primers to track phytoplasma in vectorsIdentify incubation period of X-disease phytoplasma for key vectors and relay information to growers to inform management timing intervals.Develop manuscript describing incubation period and molecular probe technique.Evaluate B. bassiana and Hirsutella sp. fungi as entomopathogenic fungiIdentify impacts of entomopathogenic fungi on incubation period using molecular probes.Develop manuscript describing effects of co-infection on pathogen incubation period.Contribute to a class on career development.Disseminate research results through two scientific conference presentations, and four grower extension talks.
Project Methods
METHODSObjective 1: Describe the X-disease infection cycle within the leafhopper vector from acquisition to transmission.Acquisition: Colladonus reductus mid-instar nymphs (15-20 days old) will be obtained from the X-disease free colony and gently brush transferred to a cage containing only phytoplasma-infected cherry trees. After a 48-h acquisition access period, nymphs will be transferred to a final cage containing uninfected host plants (sugar snap pea (Pisum sativum), and white clover (Trifolium repens)) to complete development. Presence or absence of phytoplasma will be confirmed using diagnostic PCR of plant material using X-disease specific primers (Forward: CGTTGTGGCAGCAGTTTCAG, Reverse: GAAGATGCGGTTTTCGATAGT) (Wright, Harper unpublished). All colonies and experimental cages will be maintained at 25°C, 60% relative humidity, and a16:8 (L:D) h photoperiod.Histology & Dissection: Twenty leafhoppers will be removed from the final cage at each of the following times post acquisition feeding: 7, 14, 21, 28, 35, 42, 48, and 55 days after transfer. Leafhoppers from each cohort will be anesthetized with CO2 and dissected at 32 × magnification under a dissecting microscope (Olympus SZX7, Olympus America Inc. Central Valley, PA). A Tissue Track Microscope slide (Polyscience Inc., Warrington, PA) will have four wax-barrier circles drawn onto it using an Aqua hold barrier Pap pen (Scientific Devise laboratory, Des Plaines, IL). Hemolymph will be collected and smeared into a wax-barrier circle, followed by the alimentary canal, and the primary and accessory salivary glands respectively. Slides will be air dried, then warmed to 50°C to adhere tissues to the slides. Tissue will be fixed with Carnoy's solution (Electron Microscopy Sciences, Hatfield, PA), dehydrated in ethanol, and washed in hybridization buffer. An extra 10 nymphs at each, 14, 28, and 42 days after transfer will be fixed in formalin, dehydrated in ethanol, and then infiltrated with and embedded in paraffin wax blocks. The wax blocks will be cut into 10 μm cross-sections using a Leica 2155 microtome, and will be fixed to slides by incubating at 60°C for two hours, overnight at 37°C, and then de-waxed with Histoclear II (National Diagnostics, Atlanta GA,USA). The prepared microtome sections and dissected leafhopper organs slides will be used for X-disease detection through fluorescence in situ hybridization.X-disease Phytoplasma Detection: Samples on the slides will be hybridized with high performance liquid chromatography-purified oligonucleotide probe labeled with Alexa Fluor 488 on the 5' end (Invitrogen, Carlsbad, CA) using the X-disease specific primer sequence for the probe sequence. A no probe (hybridized without fluorescent buffer) and a probe on uninfected leafhopper (clean colony) controls will be included using five leafhoppers for each control respectively. After hybridization the presence of X-disease will be examined on the microtome cross-sections and dissected organs at 200× and 400× using a fluorescence microscope with Zeiss filter set 09 (excitation wavelength = 450-490nm). Cross-sections, hemolymph, and dissected organs will be photographed using a DP74 camera mounted to the microscope to measure the change in percent fluorescence by incubation period. While the paraffin embedded cross-section approach is more time consuming than the dissections, it is complementary, as it provides intercellular location of the phytoplasma and excellent visual aids for scientific and extension publications.Data Analysis: Data will be analyzed using a logistic regression to determine the probability of X-disease phytoplasma infection among leafhopper tissue (PROC LOGISTIC, SAS Institute). We will assess the time from acquisition to being infectious by when the first salivary glands fluoresce with Phytoplasma, and determine the probability of being infectious by the number of leafhoppers with fully fluorescing salivary glands at each time point.Objective 2: Assess effects of polymicrobial infection on phytoplasma acquisition and transmission, vector fecundity and longevity.Polymicrobial effects on X-disease: Mid-instar (~3rd instar) C. reductus nymphs (15-20 days old) will be removed from the clean colony and separated into 20 groups of 10 nymphs in separate cages with non-infected developmental host plants. Groups will be randomly assigned into the following treatments: 1) B. bassiana and X-disease co-infection, 2) B. bassiana infected, 3) X-disease infected, and 4) No infection control, such that each treatment receives 5 groups (50 nymphs total). Treatments will consist of spraying the leafhopper nymphs with B. bassiana at the field rate (treatments 1 and 2) or water (treatments 3 or 4), then moving them to a cage of X-disease infected (treatments 1 and 3) or uninfected (treatments 2 and 4) cherry trees and allowing them to feed for 48 hrs before moving to a new X-disease free cage of developmental hosts. We will monitor mortality at each time point to evaluate the interactive effects of X disease phytoplasma and B. bassiana. The leafhoppers will be collected, FISH DNA probed, and examined at 200× and 400× under the fluorescent microscope. For treatment one, containing both B. bassiana and X-disease, Alexa Fluor 660 will be used to mark B. bassiana and Alexa Fluor 488 will be used to mark X-disease to allow visualization of both organisms within the cross section. B. bassiana forward primer (GAACCTACCTATCGTTGCTTC) and reverse primer (ATTCGAGGTCAACG TTCAG) are 100% specific for B. bassiana and will be used for attaching the fluorescent probes. The control treatment will also use both probes as a control for each disease.Data Analysis: From the FISH analyses we will evaluate proportional presence in the salivary glands over time using a generalized mixed linear model with a binomial (or betabinomial if overdispersed) distribution and logit link function. Leafhopper longevity in each treatment will be analyzed with a generalized linear mixed model (glmmTMB package in R) using a Gaussian, Poisson, or negative binomial distribution as appropriate, measured by AIC scores. If there are significant treatment effects, pair-wise Tukey-style comparisons will be performed to determine treatment differences in the multcomp package in R. If the three-way interaction is significant, we will use a Tukey style test to evaluate the interaction between the Beauveria presence and time, when only considering the leafhoppers exposed to X disease phytoplasma. A significant test suggests that B. bassiana alters the incubation period for X-diasease phytoplasma.EFFORTS & EVALUATIONI will present my research to fellow scientists at the Entomological Society of America (ESA) national and branch meetings. To increase transparency with the public and local growers I will present my research through 1) local grower meetings, 2) regional tree fruit extension events, and 3) my professional Instagram and Twitter accounts. I will meet bi-weekly with the full advising committee (Dr. Northfield, Cooper, and Harper) to evaluate my progress on experiment establishment for the first three months and quarterly thereafter. Once an experiment is complete, I will provide a written report to the full committee for discussion. Each objective will be considered accomplished when the research has been presented at one professional and extension conference and submitted to a peer-reviewed journal. Dr. Northfield will assess my progress of knowledge in ecological theory through my written reports and manuscript drafts. He will also gauge my mentoring and teaching abilities through meeting with my mentees, and reviewing student evaluations from the career development class. Feedback from Dr. Cooper and Dr. Harper will gauge my progress in using molecular techniques and understanding of X-disease ecology, respectively.

Progress 06/15/21 to 06/14/23

Outputs
Target Audience:Efforts during the FY 2022-2023year have been successful in reaching the target audiences planned for this grant. As outlined in the project narrative and management plan our goals were to communicate with growers through local grower meetings and regional extension events, to communicate with the advisory group of Teah Smith, Garrett Bishop, andHannah Walters, and to present research to the scientific community through two Entomological Society of America meetings. During the past year I have given 5 talks to grower and scientific communities on the vectors of X-disease and the research on tracking the phytoplasma within these vectors (10 total for granting period). Iworked bi-weekly with Teah, Garrett, and Hannah to do on farm collections of the leafhopper vectors across Washington State and to keep regional grower consulting companies aware of the most up to date information regarding the threat of X-disease transmission. I am currently registered to give a talk to a national audience of entomologist at the Entomological Society meeting in Washington DCin November. I held a workshop through WSUtraining grower consultants and scouts on the identification of leafhopper vectors and showing them the research we are doing to determine the incubation period of the phytoplasma (June, 2023). The research and indentification training was featured in the GOod Fruit Grower magazine. Changes/Problems:Throughout the project a couple challenges have arose which have caused some changes to the research. Challenge One: Our leafhopper colonies were lost over the 2021-2022 winter and needed to be restarted in summer 2022. This has delayed the start of the incubation period and polymicrobial infection trials since they require the use of non-infected mid-instar nymphs. Change/Solution: To avoid this happening again we have identified population crashes due to host plant suitability and have changed the colony plants to more preferred hosts. This year I was able to conduct transmission trials in the summer using infected branches and colony leafhoppers with a high success of acquisition. Using these methods we can now reliably infect leafhoppers with X-disease for cross-section detection and polymicrobial effects on infection.Challenge Two: Methods for tissue fixing and wax embedding leafhoppers do not exist, so I was following methods used for plant tissue wax embedding provided by Dr. Harper. When using plant tissue it is hand sliced before beginning tissue fixation to help solution penetration. Due to using whole insects and the hydrophobicity of their cuticle these methods resulted in poor fixation and wax penetration and a lack of tissue structure preservation when microtomed. Change/Solution: During the past year I have adapted the methods to enhance fixative penetration by using a 32 gauge hypodermic needle to inject the fixative, using forceps to creating openings in the cuticle, changing the amounts of time at each solution step, and using a vacuum seal to decrease pressure and enhance the wax penetration. This has greatly improved preservation of tissue structure and these methods will be published for use in other Hempiteran/leafhopper vectored plant pathogen systems. With these methods now determined combined with the high success of infecting leafhoppers, it is now possible to track the movement of phytoplasma through the leafhoppers. What opportunities for training and professional development has the project provided?This project has provided substantial opportunities for training and professional development. The research being conducted has broadened my skillset to include molecular, histological, and entompathogen research experience I did not have before. I have worked closely with scientists at the USDA forming many current and future within-discipline collaborations and working closely with Dr. Harper I have begun to form interdisciplinary collaborations with plant pathologists and horticulturalists. I have also been able to attend conferences, such as the Pacific Branch and National meetings of the Entomological Society of America, where I met and presented this research with the teams of scientists working on Peirce's Disease in California. This provided the opportunity discuss the similarities between our insect vectored plant pathogen systems and future research directions. How have the results been disseminated to communities of interest?The progress and results of the research in this project are discussed with the grower advisory group (Garrett Bishop, Teah Smith, and Hannaw Walters) every monthor as findings occur. They each represent a large portion of the stone fruit growers in Washington. Research results have also been discussed at grower meetings throughout the summer and with our WSU Tree Fruit Extension Agent (Tianna DuPont) to help disseminate results and updates to the grower community of interest as quickly and effectively as possible What do you plan to do during the next reporting period to accomplish the goals? Nothing Reported

Impacts
What was accomplished under these goals? Goal 1 - Expand into new dsciplines and pest system... Under Goal 1 substantial progress has been made for all objectives with major progress onobjectives 2 and 5 in FY 2022-2023, and major progress for objectives 1, 3, and 4 from FY 2021-2022. For objectives 2 and 4, a colony of leafhoppers has been established and a highly infected orchard was also identified in 2023. Using cut branches from the infected orchard and exposing the leafhoppers to them we have been able to determine that from a highly infected source leafhoppers can acquire X-disease after 2 days and can transmit after 20 days. We can use this as a baseline for comparison with co-infection and to get infected leafhoppers for wax embedding and probe detection to identify where the phytoplasma colonizes. For objectives 1and 3, during the past two years Dr. Harper and I have refined methods for fixing DNA and preserving leafhopper tissue structure and then embedding in wax blocks. Since this method is not common in entomological research and is novel to the leafhopper vector system we are writing a methods paper for other researchers to use in leafhopper-pathogen systems such as Xylella fastidiosa, BLTVA, and other bacteria, phytoplasma, or spiroplasma pathogens. Dr. Cooper has identified an obligate Sulcia symbiote in C. reductus which we can use as a positive control for these methods and experiments should be completed this winter. Dissections of leafhoppers have been used to confirm the microtome cross-sections cellular structures for identifying leafhopper tissue including salivary glands, alimentary canal, bacteriome, reproductive organs, and even parasitoids. These cross-sections can now be probed for X-phytoplasma and Sulcia (positive control). For objective 4, a new OMRI certified entomopathogen Beauveria bassiana product has been approved for use in cherry orchards in Washington State. I have conducteda bioassay assessing the LD50 spore concentration of the product which identified the field rate causes high mortality (90%) after 4 days and reducing the rate reduces the mortality. After this bioassay, a non-lethal concentration of the product can be used to infect leafhoppers before feeding on X-disease infected plant tissue to determine the effects of co-infection on vector competency. In collaboration with USDA researchers in Wapato, WA we are also assessing the effects infection with Beauvaria bassiana has on the insect gene regulation. This test will coincide with the experiment to determine incubation period and daily mortality to determine the probability of pathogen transmission. Goal 2: Enhance skills in molecular techniques Through this project I have gained experience in PCR and qPCR. I have extracted DNA using CTAB, amplified C. reductus leafhopper and Sulcia DNA with tested primers, and used gel electrophoresis to determine the number of C. reductus that contain Sulcia (>95%). I have also helped collect plant tissue and leafhoppers for X-disease phytoplasma detection using qPCR and became familiar with Ct score thresholds for determining if the tissue is infected or not. Fluorescent DNA probes have been created for Sulcia, and fluorescent RNA probes for X-disease phytoplasma to be tested this winter in wax cross-sections from infected leafhoppers. Goal 3: Gain experience communicating, teaching, and mentoring. The research and data collection are currently being conducted for the publications. I have assisted in mentoring 4 lab technicians, 2 master's students, and 1 Ph.D. student on career tracks, manuscript/thesis/dissertation writing, leafhopper identification, and wax embedding.

Publications


    Progress 06/15/21 to 06/14/22

    Outputs
    Target Audience:Efforts during the FY 2021-2022 year have been successful in reaching the target audiences planned for this grant. As outlined in the project narrative and management plan our goals were to communicate with growers through local grower meetings and regional extension events, to communicate with the advisory group of Teah Smith, Garrett Bishop, and Hannah Walters, and to present research to the scientific community through two Entomological Society of America meetings. During the past year I have given 5 talks to grower and scientific communities on the vectors of X-disease and the research on tracking the phytoplasma within these vectors. I have worked weekly with Teah, Garrett, and Hannah to do on farm collections of the leafhopper vectors across Washington State and to keep regional grower consulting companies aware of the most up to date information regarding the threat of X-disease transmission. I am currently registered to give a talk to an international audience of entomologist at the Entomological Society meeting in Vancouver, BC Canada in November. I have held a workshop with Wilbur-Ellis consultants training them on the identification of leafhopper vectors and showing them the research we are doing to determine the incubation period of the phytoplasma. Changes/Problems:Over the past year challenges have arose which have caused some changes to the research. Challenge One: Our leafhopper colonies were lost over the 2021-2022 winter and needed to be restarted this Spring. This has delayed the start of the incubation period and polymicrobial infection trials since they require the use of non-infected mid-instar nymphs. Change/Solution: To avoid this happening again the Northfield lab now has 3 Percival temperature and humidity controlled chambers for keeping the colonies. This year I have been collecting and testing the wild population of leafhoppers for X-disease and found that <5% of adults in highly X-infected cherry blocks have the phytoplasma in detectable quanitites so as a back up we can use the collection of wild nymphs for these experiments next year (2023). Challenge Two: Methods for tissue fixing and wax embedding leafhoppers do not exist, so I was following methods used for plant tissue wax embedding provided by Dr. Harper. When using plant tissue it is hand sliced before beginning tissue fixation to help solution penetration. Due to using whole insects and the hydrophobicity of their cuticle these methods resulted in poor fixation and wax penetration and a lack of tissue structure preservation when microtomed. Change/Solution: During the past year I have adapted the methods to enhance fixative penetration by using a 32 gauge hypodermic needle to inject the fixative, using forceps to creating openings in the cuticle, changing the amounts of time at each solution step, and using a vacuum seal to decrease pressure and enhance the wax penetration. This has greatly improved preservation of tissue structure and these methods will be published for use in other Hempiteran/leafhopper vectored plant pathogen systems. What opportunities for training and professional development has the project provided?This project has provided substantial opportunities for training and professional development. The research being conducted has broadened my skillset to include molecular, histological, and entompathogen research experience I did not have before. I have worked closely with scientists at the USDA forming many current and future within-discipline collaborations and working closely with Dr. Harper I have begun to form interdisciplinary collaborations with plant pathologists and horticulturalists. I have also been able to attend conferences, such as the Pacific Branch Entomological Society of America meeting, where I met and presented this research with the teams of scientists working on Peirce's Disease in California. This provided the opportunity discuss the similarities between our insect vectored plant pathogen systems and future research directions. How have the results been disseminated to communities of interest?The progress and results of the research in this project are discussed with the grower advisory group (Garrett Bishop, Teah Smith, and Hannaw Walters) every other week or as findings occur. They each represent a large portion of the stone fruit growers in Washington. Research results have also been discussed at grower meetings throughout the summer and with our WSU Tree Fruit Extension Agent (Tianna DuPont) to help disseminate results and updates to the grower community of interest as quickly and effectively as possible. What do you plan to do during the next reporting period to accomplish the goals?During the next reporting period (2022-2023) I plan to begin using the histological methods that we have refined during this past year to determine the pathway of X-disease and its incubation period within the leafhopper vectors. I will also continue to determine the direct (mortality) and indirect effects of B. bassiana on the leafhoppers and their ability to acquire and transmit X-disease. I will attend the national (Fall 2022) and branch meeting (Spring 2023) of the Entomological Society of America to report my findings to the scientific community and share methods for those working on similar insect-pathogen systems. I will also attend grower meetings and the Orchard Pest and Disease Management Conference this fall to disseminate the results to the target grower and industry stakeholders. After the experiments are concluded I will write a methods paper for the histological cross-sectioning and fluorescent probe pathogen identification and a paper on the effects of B. bassiana on X-disease vectors and their ability to transmit the phytoplasma.

    Impacts
    What was accomplished under these goals? Goal 1 - Expand into new dsciplines and pest system... Under Goal 1 substantial progress has been made for objectives 1, 2, 3, and 4. During the past year Dr. Harper and I have refined methods for fixing DNA and preserving leafhopper tissue structure and then embedding in wax blocks. Since this method is not common in entomological research and is novel to the leafhopper vector system we are writing a methods paper for other researchers to use in leafhopper-pathogen systems such as Xylella fastidiosa, BLTVA, and other bacteria, phytoplasma, or spiroplasma pathogens. Dr. Cooper has identified an obligate Sulcia symbiote in C. reductus which we can use as a positive control for these methods and experiments should be completed this fall. Dissections of leafhoppers have been used to confirm the microtome cross-sections cellular structures for identifying leafhopper tissue including salivary glands, alimentary canal, bacteriome, and reproductive organs. These cross-sections can now be probed for X-phytoplasma and Sulcia (positive control). For objective 4, a new OMRI certified Beauveria bassiana product has been approved for use in Cherry orchards in Washington State this summer. I have begun a bioassay assessing the LD50 spore concentration of the product which is currently ongoing (Day 14). After this bioassay, a non-lethal concentration of the product will be used to infect leafhoppers before feeding on X-disease infected plant tissue to determine the effects of co-infection on vector competency. This test will coincide with the experiment to determine incubation period and daily mortality to determine the probability of pathogen transmission. Goal 2: Enhance skills in molecular techniques Through this project I have gained experience in PCR and qPCR. I have extracted DNA using CTAB, amplified C. reductus leafhopper and Sulcia DNA with tested primers, and used gel electrophoresis to determine the number of C. reductus that contain Sulcia (>95%). I have also helped collect plant tissue and leafhoppers for X-disease phytoplasma detection using qPCR and become familiar with CT score thresholds for determining if the tissue is infected or not. Fluorescent DNA probes have been created for Sulcia, and fluorescent RNA probes for X-disease phytoplasma to be tested this fall in wax cross-sections. Goal 3: Gain experience communicating, teaching, and mentoring. The research and data collection are currently being conducted for the publications. I have assisted in mentoring 2 lab technicians, 2 master's students, and 1 Ph.D. student on career tracks, manuscript/thesis/dissertation writing, leafhopper identification, and wax embedding. I plan to lead a University Seminar discussionon Career Development by Spring 2023.

    Publications