Source: UNIVERSITY OF CALIFORNIA, RIVERSIDE submitted to
ENVIRONMENTAL DYNAMICS OF ECOSYSTEM ENGINEERING ANTS AND THEIR MODIFICATIONS OF SOIL STRUCTURE
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
1022904
Grant No.
2020-67034-31891
Cumulative Award Amt.
$70,344.00
Proposal No.
2019-07320
Multistate No.
(N/A)
Project Start Date
Jul 1, 2020
Project End Date
Jun 30, 2023
Grant Year
2020
Program Code
[A7101]- AFRI Predoctoral Fellowships
Recipient Organization
UNIVERSITY OF CALIFORNIA, RIVERSIDE
(N/A)
RIVERSIDE,CA 92521
Performing Department
Entomology
Non Technical Summary
As soil ecosystem engineers, ants contribute to the maintenance of soil fertility, enriching and aerating soil through foraging and nest excavation. Through this ecosystem service, ants facilitate the development of vegetation and microorganism communities, a vital aspect of agricultural production. However, the impact of ants as soil engineers is likely to differ across climates, and their ecosystem services may be vulnerable under climate change. The goal of this predoctoral research project is to reveal how landscapes and environments affect soil movement by ants and predict the vulnerability of this ecosystem service under climate change. Specifically, I seek to understand the effects of temperature, soil type, and vegetation on nest structure and soil movement by ants, and utilize genomic data to predict distributional shifts under climate projections, of two species: Formica aerata, prominent throughout California's agricultural land, and Formica podzolica, common in North American forests. First, I will determine how ant-soil interactions change with environmental factors by measuring their nest shape and soil movement in the lab under various temperatures and using multiple soil types and plants. Second, I will reveal how these species are adapted to climatic heterogeneity through genomic sequencing. Third, I will predict geographic shifts in the ants' soil services by modeling the environmental conditions that will best allow these ant species to survive under future climate change scenarios. Through these methods, this project hopes to achieve the ultimate goal of better-informing scientists and growers about how ants contribute to soil ecosystem maintenance and how to manage them best to optimize soil fertility.
Animal Health Component
10%
Research Effort Categories
Basic
90%
Applied
10%
Developmental
(N/A)
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
1013110113050%
2113110113050%
Goals / Objectives
The goal of this predoctoral research project is to reveal how landscapes and climates affect soil movement by ants and predict the vulnerability of this ecosystem service under climate change. Specifically, I seek to understand the effects of temperature, soil type, and vegetation on nest structure and soil movement by ants, and utilize genomic data to predict distributional shifts under climate projections, of two species: Formica aerata, prominent throughout California's agricultural land, and Formica podzolica, common in North American forests. ObjectivesDetermine how ant-soil interactions change with environmental factors using transplant experiments. Reveal how these species are adapted to climatic heterogeneity through genomic markers. Predict geographic shifts in the ants' soil services using environmental niche modeling.
Project Methods
Objective 1. I will collect 100 workers from each of 20 colonies per species, from both low and high elevations, house them in transparent nest boxes (18"x20"x3/8'') in a laboratory setting, and allow them one week to excavate nests. For F. aerata, the low-elevation sites will be orchards in Riverside, CA, and the high-elevation sites will be orchards in Julian, CA. For F. podzolica, the low-elevation sites will be at the foothills of the Rocky Mountains in Boulder, CO, and the high-elevation sites will be in the subalpine zone around Ward, CO. I will provide the ants with water and feed them a diet of sugar water. I will test treatments of temperature, soil type, and presence/absence of vegetation in a progressive manner throughout three sub-experiments. First, I will examine the effects of temperature on bioturbation and nest structure, applying heat to half the boxes via temperature chambers that envelop the soil surface, and applying a consistent, but cooler, temperature via similar temperature chambers to the other half as controls. Second, I will compare sandy and loamy soil types using the temperature treatment that produces the largest effect size from the first experiment. Third, I will compare the presence and absence of vegetation using whichever soil/temperature combination produces the largest effect size from the second experiment. Each of these experiments will be replicated twice, for a total of 40 colony replications per species for each experiment.I will keep the nest boxes in a room at 60 °F to mimic a subterranean nest temperature. The glass walls of the boxes will be kept dark with black plastic to mimic an underground habitat. Four temperature chambers will encapsulate five boxes each from the soil surface up, with the majority of the area of the nest boxes extending below the chamber and in contact with the ambient air temperature. I have already built these boxes and chambers, and they are currently undergoing pilot testing with F. podzolica, which are excavating significant tunnels. One chamber will have the high-temperature treatment that will mimic the average summer temperature of the low-elevation sites, and the other chamber will have the low-temperature treatment that will mimic the average summer temperature of the high-elevation sites. The heat will be produced by a thermal mass consisting of a concrete mortar mix surrounding radiant heating tubing, which will connect to a temperature-controlling outlet. I will monitor soil temperature using iButtons at various depths. Each nest box will have a treatment of either loamy or sandy soil, and I will add a thin layer of colored sand every 10 cm (a different color for every layer). The colors will allow me to assess the movement of the soil layers visually. For the vegetation treatment, I will plant one apple sapling (a crop around which F. aerata has been found living) in each nest box, standardizing by above- and below-ground biomass. I have already built these boxes and chambers, and they are undergoing pilot testing. During the excavation period, I will take photographs daily to measure two attributes of the subterranean nest excavation. First, I will quantify bioturbation using the layers of soil denoted by different colors of sand. I will overlay a grid on each photo using Adobe Illustrator and measure the quantity and distance of colored sediment moved by the ants from their respective horizons within the box. This data will illuminate how environmental factors influence the extent of soil mixing by these ants. Second, I will quantify the nest structure by measuring the dimensions (length and depth) and the total number and depth of chambers at experiment completion. This data will inform us about the extent to which ants can aerate soil under the influence of critical environmental factors. I will perform all statistical analyses in R. I will use the lme4and lmerTestpackages to build multiple linear mixed-effects models. In these models, original colony elevation, temperature treatment, soil type, and presence/absence of vegetation will be fixed effects, and time will be a random effect. Distance and direction of displaced soil, in addition to tunnel number, depth, and dimensions, will be the response variables.Objective 2. Knowledge about climate-associated genomic adaptations, paired with empirical data about ant nest adaptations from Objective 1, will allow for a multifaceted understanding of the vulnerability of each species under different climate and landscape scenarios. For F. podzolica, I will use samples collected across the entire range during 2016-2019 by the Purcell and Brelsford Labs at UCR. For F. aerata, I will sample across California and Oregon during fall 2021. To prepare samples for sequencing, I will conduct DNA tagmentation using TDE1 enzyme, ligate barcoded adapters, and remove small DNA fragments.I will sequence whole genomes on the Illumina HiSeq X platform at the UCR Institute for Integrative Genome Biology Genomics Core.To determine the relative contribution of environmental versus geographic variation to overall genomic variation, I will use a Mantel test and follow up with a multiple regression analysis. For these analyses, I will use WorldClimglobal climate data and soil type data from the USDA National Cooperative Soil Survey. I will conduct a principal components analysis, which will allow me to identify the environmental variables that differ most between populations, and then perform a gradient forest analysis (GFA), which will rank the ecological variables correlated with genomic variation. Finally, I will determine which genomic regions are most affected by environmental factors by conducting an environmental association analysis. In this analysis, I will correlate SNPs with top-ranking ecological variables using a latent factor mixed model and search for outlier SNPs using the Bayenv2 method.Using climate projections from global climate models used in the Fifth Assessment Intergovernmental Panel on Climate Change (IPCC) Report,I will extend the GFA to investigate which populations might be most vulnerable to climate change based on current and predicted genomic variation.Objective 3. I will employ environmental niche modeling methods to predict how the species ranges will shift based on their calculated genomic vulnerability. I will produce a climate envelope model for the future affected species distributions and ecosystem services, altering climate variables (those I identified to correlate with genomic variation) across the species' ranges according to the IPCC predictions (i.e. shifting the climate space) and subsequently map the predicted climate affected distributions (CAD) of species back into geographical space. I will assess changes in distribution patterns (and, therefore, the soil ecosystem services associated with the species) owing to predicted climate change by comparing the current distribution with the CAD. Finally, I will predict soil aeration and bioturbation in regions across the species ranges, informed by Objective 1.Efforts: I will engage the general public and stakeholdersin the results of this research by holding an outreach booth at the annual Riverside Insect Fair and collaborating with the extension specialists in the UCR Entomology department to disseminate the research findingsEvaluation: Quantitative data to be collected: nest architecture and bioturbation data andant genomic data for each species. Models to be produced: landscape genomics models for each ant species, environmental niche models for each ant species.

Progress 07/01/20 to 06/30/23

Outputs
Target Audience:During my research project, I strategically focused on engaging diverse target audiences, each of whom played a crucial role in advancing my work and its impact. These audiences include fellow scientists, undergraduate and graduate students, extension faculty, stakeholders (such as growers), federal funders, agricultural industries, and the public. 1.Fellow scientists: Fellow researchers and experts in entomology constitute a primary audience. My research matters to them because it contributes to the body of scientific knowledge, and the findings may have significant implications for their work. I reached this audience through conference presentations, where I shared my findings, methodologies, and insights with my peers. Additionally, publishing my work in scientific journals allowed for in-depth examination and peer review, further cementing my work's relevance within the scientific community. 2.Undergraduate students: Educating the next generation of scientists is vital. My work has mattered to students because it offered them opportunities to learn and gain hands-on experience in entomology. I engaged students through laboratory instruction, providing valuable practical knowledge and fostering a passion for research. 3.Extension faculty: Extension faculty play a critical role in bridging the gap between research and practical applications in the agriculture industry. My research matters to them because it can inform their recommendations and outreach to growers and farmers. I reached extension faculty through knowledge-sharing sessions tailored to their needs and interests. 4.Stakeholders (growers): Growers and agricultural stakeholders are deeply invested in the outcomes of my research. My work can directly impact their practices, crop yields, and overall economic viability. Engaging with stakeholders involved listening to their concerns and ensuring that my research addressed their practical challenges. 5.General public: My research also matters because it can have broader implications for food security and environmental sustainability. I engaged the public through outreach efforts, which included publishing articles in popular science outlets. These articles helped to disseminate my findings and their broader significance to a non-specialized audience. In summary, the comprehensive approach to targeting these diverse audiences demonstrates my commitment to making a meaningful impact with my research. By tailoring my engagement strategies to each group's specific needs and interests, I have effectively communicated the relevance of my work and its potential to address a range of scientific, economic, and societal challenges. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?Under the guidance of my mentor, Jessica Purcell, I received personalized mentorship encompassing various aspects of scientific research. This mentorship covered experimental design, statistical analysis, and writing scientific research articles. Furthermore, I had the privilege of presenting the outcomes of my work at conferences. Notably, my research has significantly impacted students by providing them with valuable opportunities to delve into the world of entomology. I actively engaged students through hands-on laboratory instruction, equipping them with practical knowledge and nurturing a genuine passion for research. How have the results been disseminated to communities of interest?I connected with various audiences effectively. At conferences, I shared my findings, methodologies, and insights with fellow scientists, strengthening the bonds within our research community. The publication of my work in scientific journals allowed for a more profound examination and peer review, solidifying its relevance in the scientific realm. I held knowledge-sharing sessions with extension faculty to address their needs and interests. This approach ensured that the information I provided aligned closely with their goals. In my interactions with growers, I made a point to actively listen to their concerns, guaranteeing that my research directly addressed the practical challenges they faced. To engage the public, I undertook outreach efforts, including publishing articles in popular science outlets. These articles served as a bridge, conveying my findings and their broader significance to a wider, non-specialized audience, promoting a deeper understanding of my research's real-world relevance. What do you plan to do during the next reporting period to accomplish the goals? Nothing Reported

Impacts
What was accomplished under these goals? 1. Determine how ant-soil interactions change with environmental factors using transplant experiments. In a full-factorial experiment, I investigated the effects of two temperature treatments on the shape and size of nests built by Formica podzolica ants collected from high and low elevations in the Colorado Rocky Mountains. Ants nested in experimental chambers with soil surface temperatures matching the local temperatures of sample sites. I observed a plastic response of nest architecture to conditions experienced during excavation; workers experiencing a high temperature excavated deeper nests than those experiencing a cooler temperature. Further, I found evidence of local adaptation to temperature, with a significant interaction effect of natal elevation and temperature treatment on nest size and complexity. Specifically, workers from high-elevation sites built larger nests with more tunnels when placed in the cool surface temperature treatment, and workers from low-elevation sites exhibited the opposite pattern. My results suggest that subterranean ant nest architecture is shaped by a combination of plastic and locally adapted building behaviors; I suggest that the flexibility of this 'extended phenotype' likely contributes to the widespread success of ants. My research on these hardworking ants has benefits for both growers and our society as a whole. For growers, understanding how ants build their nests, especially in response to temperature variations, can help improve farming practices. Ants play a vital role in ecosystems by providing services like soil aeration and nutrient cycling. We can potentially apply my insights on how ants adapt their nest-building to different temperatures to optimize soil health and crop growth. This research may lead to better pest control and more productive agricultural systems, benefiting farmers and food production. For society, studying ants' underground homes offers a fascinating glimpse into the intricate ways nature adapts to its surroundings. It shows us that these tiny creatures are not just responding to their environment but also shaping it. By learning how ants adjust their nest architecture to local conditions, we gain a deeper appreciation for the complexity of life on Earth. This knowledge contributes to our understanding of ecological processes and biodiversity. It underscores how interconnected all living organisms are and highlights the remarkable adaptability of these insects, which has allowed them to thrive in various environments. This research broadens our scientific horizons and enhances our appreciation of the natural world. 2.Reveal how these species are adapted to climatic heterogeneity through genomic markers. I conducted genomic analyses to identify markers associated with climatic heterogeneity in a widespread species, F. podzolica. Genomic signatures of adaptation to temperature, precipitation, and seasonality were present across F. podzolica's range, with one locus exhibiting a precipitation-associated alternative allele exclusively at the northern edge of the range. This study suggests that genomic variation may factor in adaptive potential, especially at range margins. This research on F. podzolica ants has important implications for both growers and society as a whole. Understanding how these ants adapt to different climates can be incredibly valuable for growers. These tiny creatures influence soil properties, and by identifying the genetic markers linked to their adaptation to temperature, precipitation, and seasonal changes, we can gain insights into how to manage soil conditions effectively. This knowledge may help farmers optimize their agricultural practices, producing more resilient and productive crops. It could contribute to sustainable farming methods that can withstand the challenges of changing weather patterns and, ultimately, benefit food production. For society, this research reveals the incredible adaptability and diversity of life on our planet. It shows us that even the smallest organisms, like ants, can adjust to different climates, and this adaptability is often linked to their genetic makeup. Understanding how these ants cope with varying environmental conditions gives us a deeper appreciation for the intricate web of life surrounding us. This research enriches our scientific knowledge and reminds us of how nature evolves and thrives. It also highlights the importance of preserving biodiversity and the delicate balance of ecosystems in the face of climate change. Ultimately, this work contributes to our broader understanding of the natural world and the importance of protecting it for future generations. 3. Predict geographic shifts in the ants' soil services using environmental niche modeling. Environmental niche modeling for this goal is currently underway.

Publications

  • Type: Theses/Dissertations Status: Accepted Year Published: 2022 Citation: Sankovitz, M. (2022). An Interdisciplinary Approach to Understanding Environmental Dynamics of Soil Ecosystem Engineering Ants. University of California, Riverside.


Progress 07/01/21 to 06/30/22

Outputs
Target Audience:The target audience reached during this reporting period was entomologists and growers working with ants. I gave a presentation, wrote multiple blog posts explaining my research topic, and provided guidance for working with ants. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?The PD received one-on-one mentorship from advisor Jessica Purcell in experimental design, statistical analysis, and the writing of a scientific research article. Additionally, the PD presented the preliminary results of Objective 2 in a talk at the Entomology 2021 conference. How have the results been disseminated to communities of interest?The results of Objective 1 were published in Scientific Reports (Sankovitz & Purcell 2021), and the preliminary results of Objective 2 were presented at the Entomology 2021 conference. What do you plan to do during the next reporting period to accomplish the goals?To accomplish Objectives 2 and 3, I will continue to use landscape genomics methods to identify genomic signals of adaptation across two ant species ranges, which will allow me to estimate the vulnerability of populations under changing climates. For F. podzolica, I will use samples collected across the entire range during 2016-2019 by the Purcell and Brelsford Labs at UCR. For F. aerata, I will sample across California and Oregon this fall. To prepare samples for sequencing, I will conduct DNA tagmentation using TDE1 enzyme, ligate barcoded adapters, and remove small DNA fragments. Then, I will sequence whole genomes on the Illumina HiSeq X platform at the UCR Institute for Integrative Genome Biology Genomics Core. To determine the relative contribution of environmental versus geographic variation to overall genomic variation, I will use a Mantel test and follow up with a multiple regression analysis. I will use WorldClim global climate data and soil type data from the USDA National Cooperative Soil Survey for these analyses. Next, I will conduct a principal components analysis, which will allow me to identify the environmental variables that differ most between populations, and then conduct a gradient forest analysis (GFA), which will rank the environmental variables correlated with genomic variation. Finally, I will determine which genomic regions are most affected by environmental factors by conducting an environmental association analysis. I will correlate SNPs with top-ranking environmental variables using a latent factor mixed model and search for outlier SNPs using the Bayenv2 method. Using climate projections from global climate models used in the Fifth Assessment Intergovernmental Panel on Climate Change (IPCC) Report, I will extend the GFA to investigate which populations might be most vulnerable to climate change based on current and predicted genomic variation.

Impacts
What was accomplished under these goals? Social insects are among the most abundant arthropods in terrestrial ecosystems, where they provide essential ecosystem services. For instance, subterranean ants help maintain healthy soil through aeration, decomposition, and nutrient cycling. The effect of ant colonies on soil properties has been studied for decades, yet little is known about ant nest properties, like architecture, due to the difficulty of observing these belowground patterns in most species. Furthermore, many ant species' ranges span environmental gradients, and their nest architecture and nest-building behavior are likely shaped by the climatic and landscape features of their specific habitats. However, we do not know how climate plays a role in ants' ability to be ecosystem engineers and how this might shift with climate change. During this reporting period, we worked toward Objectives 1 and 2 by exploring how temperature shapes an 'extended phenotype' of ant colonies - nest architecture - and investigating genomic signatures of local adaptation to climate. Our results from Objective 1 suggest that subterranean ant nest architecture is shaped by a combination of plastic and locally adapted building behaviors; we suggest that the flexibility of this extended phenotype likely contributes to the widespread success of ants. We published this study in Scientific Reports (Sankovitz & Purcell 2021). For Objective 2, we sequenced whole genomes of 152 Formica podzolica ants sampled from across North America. We are currently analyzing the genomic data alongside remote-sensed climate data to find regions of the genome that have candidate genes for climate adaptation. This information will allow us to understand which climate variables have the most significant impact on these ants and predict future population shifts under climate change. This work, completed under the mentorship of Jessica Purcell (University of California Riverside), has resulted in new fundamental knowledge significant enough to be included in a publication, novel methods and techniques, and improved skills for the PD. We are in the process of developing a predictive understanding of ants' impact on soil. This work aligns with the AFRI Farm Bill priority areas "Bioenergy, Natural Resources, and Environment" and "Plant Health and Production and Plant Products". In addition, it is consistent with the goals of AFRI EWD by preparing me via interactive mentoring, scientific training, career development, and self-evaluation.

Publications

  • Type: Journal Articles Status: Published Year Published: 2021 Citation: Sankovitz, M., & Purcell, J. (2021). Ant nest architecture is shaped by local adaptation and plastic response to temperature. Scientific reports, 11(1), 1-10.


Progress 07/01/20 to 06/30/21

Outputs
Target Audience:The target audience reached during this reporting period was entomologists and growers working with ants. I gave remote presentations, wrote multiple blog posts explaining my research topic, and provided guidance for working with ants. Changes/Problems:In Objective 1, I had planned on carrying outsoil-type and vegetation experiments. I introduced colonies to either sandy or loamy soil but could only carry out one trial of 20 colonies because the ants were unable to build nests in the sandy soil. The nest boxes ended up being too small to reasonably contain plants, and the field season also did not contain enough time to carry out this experiment. To examine bioturbation, I added a thin layer of colored sand every 10 cm (a different color for every layer) to one trial of 20 colonies (similar to Halfen & Hasiotis 2010). The aim of these colors was to allow me to assess the movement of the soil layers visually, but the ants uniformly stopped digging once they reached the first layer of sand. This was unexpected, as the harvester ants used in Halfen & Hasiotis 2010 were not sensitive to layers of various substrates. I excluded these sand layers from the rest of the experimental trials of nest architecture so as not to influence the ways the ants built their tunnels. What opportunities for training and professional development has the project provided?The PD received one-on-one mentorship from advisor Jessica Purcell in experimental design, statistical analysis, and the writing of a scientific research article. Additionally, the PD mentored three undergraduatesduring the data collection and analysis phases of this reporting period. Finally, the PD presented the results of this research in a talk at the Entomology 2020 conference. How have the results been disseminated to communities of interest? Nothing Reported What do you plan to do during the next reporting period to accomplish the goals?To accomplish Objectives 2 and 3, I willutilize landscape genomics methods to identify genomic signals of adaptation across two ant species ranges, which will allow me to estimate the vulnerability of populations under changing climates. For F. podzolica, I will use samples collected across the entire range during 2016-2019 by the Purcell and Brelsford Labs at UCR. For F. aerata, I will sample across California and Oregon this fall. To prepare samples for sequencing, I will conduct DNA tagmentation using TDE1 enzyme, ligate barcoded adapters, and remove small DNA fragments. Then, I will sequence whole genomes on the Illumina HiSeq X platform at the UCR Institute for Integrative Genome Biology Genomics Core. To determine the relative contribution of environmental versus geographic variation to overall genomic variation, I will use a Mantel test and follow up with a multiple regression analysis. I will use WorldClimglobal climate data and soil type data from the USDA National Cooperative Soil Survey for these analyses. Next, I will conduct a principal components analysis, which will allow me to identify the environmental variables that differ most between populations, and then conduct a gradient forest analysis (GFA), which will rank the environmental variables correlated with genomic variation. Finally, I will determine which genomic regions are most affected by environmental factors by conducting an environmental association analysis. I will correlate SNPs with top-ranking environmental variables using a latent factor mixed model and search for outlier SNPs using the Bayenv2 method.Using climate projections from global climate models used in the Fifth Assessment Intergovernmental Panel on Climate Change (IPCC) Report.I will extend the GFA to investigate which populations might be most vulnerable to climate change based on current and predicted genomic variation.

Impacts
What was accomplished under these goals? Social insects are among the most abundant arthropods in terrestrial ecosystems, where they provide essential ecosystem services. For instance, subterranean ants help maintain healthy soil through aeration, decomposition, and nutrient cycling. The effect of ant colonies on soil properties has been studied for decades, yet little is known about ant nest properties, like architecture, due to the difficulty of observing these belowground patterns in most species. Furthermore, many ant species' ranges span environmental gradients, and their nest architecture and nest-building behavior are likely shaped by the climatic and landscape features of their specific habitats. During this reporting period, we worked toward Objective 1 by exploringhow temperature shapes an 'extended phenotype' of ant colonies - nest architecture. Using a full factorial experiment, we investigated the nest architecture of montane Formica podzolica ant workers collected from high and low elevations in the Colorado Rocky Mountains. We allowed ants to nest in experimental chambers with soil surface temperatures matching the approximate local temperatures of high and low sample sites and measured nest architecture characteristics. We observed a plastic response of nest architecture to conditions experienced during nest construction; workers experiencing a high temperature excavated deeper nests than those experiencing a cooler temperature. Further, we found evidence of local adaptation to the different temperature conditions, with a significant interaction effect of natal elevational and temperature treatment on nest size and complexity. Our results suggest that subterranean ant nest architecture is shaped by a combination of plastic and locally adapted building behaviors; we suggest that the flexibility of this extended phenotype likely contributes to the widespread success of ants. This work, completed under the mentorship of Jessica Purcell (University of California Riverside), has resulted in new fundamental knowledge significant enough to be included in a publication, novel methods and techniques, and improved skills for the PD. We are in the process of developing a predictive understanding of ants' impact on soil and aligns with the AFRI Farm Bill priority areas "Bioenergy, Natural Resources, and Environment" and "Plant Health and Production and Plant Products". In addition, it is consistent with the goals of AFRI EWD by preparing me via interactive mentoring, scientific training, career development, and self-evaluation.

Publications