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Phosphorus (P) is one of the three major macronutrients that plants need to grow and survive. When P is scarce, plants utilize a network of characterized responses known as the Phosphate Starvation Response (PSR) to remobilize internal stores of P as well as external P from soil. Emerging evidence shows the PSR is regulated by a specialized group of secondary messenger molecules, inositol pyrophosphates (PP-InsP). PP-InsPs and their precursors, inositol phosphates (InsPs), are important for plant abiotic stress responses, hormone signaling, and other stress responses. While PP-InsPs are critical for plant survival, much about the roles of PP-InsPs and how they are regulated remains to be understood. Further, the enzymes responsible for the synthesis of PP-InsPs in plants have been recently discovered; however, not much is known about the enzymes that degrade PP-InsPs in plants. The goal of the work presented herein is to understand critical aspects of the PP-InsP signaling in plants and leverage this information into a P phytoremediation strategy. To achieve this, I have investigated a group of PP-InsP phosphatases and assessed long-term impacts of depleting PP-InsPs in two plant species, Arabidopsis thaliana (Arabidopsis) and Thlaspi arvense (Pennycress). Exploring the impact of plant PP-InsP phosphatases has allowed me to explore critical aspects of PP-InsP sensing that show great promise for informing P remediation strategies. / Doctor of Philosophy / The Phosphorus (P) crisis presents a major challenge to food security. While Phosphorus (P) is critical for crop growth, P is a nonrenewable and increasingly limited resource. Our global population is fed at the expense of the remaining mineable P reserve, which may be depleted in as early as 30 years. Further, fertilizer runoff from farmland and urban areas poses a dangerous problem as increased nutrients in watersheds toxifies our water supply and aquatic ecosystems. Time is running out to preserve our P supply. New and innovative strategies that reduce fertilizer inputs and watershed pollution are key to securing the global food supply and protecting the environment. Emerging evidence shows plants sense and respond to P using signaling molecules known as inositol pyrophosphates (PP-InsPs). My work and that of others are key in showing that alteration of the levels of PP-InsPs can decrease plant P dependency or cause plants to hyperaccumulate P. Understanding how plants are able to sense, respond, and acquire P is crucial to inform future P phytoremediation strategies to secure global food security.

Phosphate (Pi) is an essential nutrient for plants, required for plant growth and seed viability. When Pi is limited, plants undergo dynamic morphological and metabolic changes to leverage available Pi, known as the Phosphate Starvation Response (PSR). The inositol phosphate (InsP) signaling pathway is a crucial element of the plant's ability to regulate the PSR and respond to changing energy conditions. InsPs are synthesized from the cyclic 6-carbon polyol scaffold, myo-inositol. Inositol hexakisphosphate (InsP6) is the most abundant InsP signaling molecule and can be phosphorylated by the multifunctional inositol tetrakisphosphate 1-kinase 1 (ITPK1) and diphosphoinositol pentakisphosphate (VIP) kinases, resulting in inositol pyrophosphates (PP-InsPs). PP-InsPs have high energy bonds and have been linked to Pi maintenance and energy homeostasis in yeast, plants, and mammals. However, the precise mechanism(s) by which PP-InsPs act within plant signaling pathways remains to be determined. Two approaches to understand the role of PP-InsPs in plants are described within this dissertation. The first approach analyzes genetic loss-of-function vip1/vip2 double mutants, and their responses to low Pi conditions. Specifically, vip1/vip2 double mutant gene expression and lipid remodeling patterns in response to low Pi were characterized. We found that vip1-2/vip2-2 had an impacted lipid remodeling response under low Pi conditions, whereas ipk1 had altered lipid composition under Pi-replete conditions. In a complementary approach, a gain-of-function in either the ITPK1 or the kinase domain of VIP (VIP2KD) were constructed in transgenic Arabidopsis thaliana plants. Both ITPK1 and VIP2KD transgenic plants contain elevated levels of the specific inositol pyrophosphate, InsP8. Elevated InsP8 in both types of plants results in changes in growth and senescence phenotypes, delayed time to flowering, Pi accumulation, and altered PSR gene expression. The data from both approaches suggest new roles for PP-InsPs in the regulation of the PSR and other signaling pathways in plants. To enhance my teaching and leadership skills, I participated in the Graduate Teaching Scholars (GTS) program. As a GTS, I worked with the Virginia Tech Research and Extension Experiential Learning (VT-REEL) program where I developed a structured mentorship program for undergraduate and graduate students and created a professional development workshop series. During the COVID-19 pandemic, I developed an online version of the VT-REEL program. Using inclusive pedagogy practices and surveys from the participants, we compiled the best practices for moving a summer undergraduate research program online. These practices come from surveyed participants in the 2020 and provides strategies that can be tailored to various online research experiences and be implemented in both online and in-person formats. / Doctor of Philosophy / Phosphate (Pi) is crucial for plant development and crop yield, but is often limited in soils. Pi-containing fertilizers are often added to supplement soils. Overuse of Pi-containing fertilizers can lead to Pi runoff and can devastate aquatic ecosystems. In addition, Pi is a limited, nonrenewable resource, with U.S. stores projected to be depleted in as little as 30 years. It is now crucial to develop crops that can feed a growing population with less Pi input. Here, we describe how changing levels of plant messenger molecules known as inositol pyrophosphates (PP-InsPs) impact the ability of plants to sense and respond to Pi. This knowledge advances understanding f how mineral nutrient physiology affects many plants traits, and can be harnessed to develop novel strategies to reduce Pi-application and overuse.