To maintain cellular homeostasis, concentrations, chemical speciation, and localization of mineral nutrients and toxic trace elements need to be regulated. Imaging the cellular and subcellular localization of elements and measuring their in situ chemical speciation are challenging tasks that can be undertaken using synchrotronbased techniques, such as X-ray fluorescence and X-ray absorption spectrometry, and mass spectrometry-based techniques, such as secondary ion mass spectrometry and laser-ablation inductively coupled plasma mass spectrometry. We review the advantages and limitations of these techniques, and discuss examples of their applications, which have revealed highly heterogeneous distribution patterns of elements in different cell types, often varying in chemical speciation. Combining these techniques with molecular genetic approaches can unravel functions of genes involved in element homeostasis.
Spatial and chemical information of mineral element homeostasisPlants take up a range of mineral elements from the soil, some of which are essential for growth, whereas others are non-essential [1]. Deficiencies of essential elements are a major limiting factor for crop production in many areas worldwide, whereas excessive accumulation of both essential and non-essential elements can lead to phytotoxicity [1]. Accumulation of some elements, such as cadmium (Cd) [2] and arsenic (As) [3], in the edible parts of crops may pose a significant risk to human health well before phytotoxicity occurs. By contrast, there is a need to increase essential micronutrients, such as iron (Fe) and zinc (Zn), in plantbased foods to alleviate their deficiencies in humans [4]. Plant nutrition research aims to understand how minerals are acquired, transported, distributed, stored, and used in plants. This knowledge is important not only for sustainable agricultural production but also for ensuring the nutritional quality and safety of agricultural products [5].Analyses of total elemental concentrations can now be performed using high-throughput platforms to reveal the ionomic profile (see Glossary) of plant tissues [6]. Although the total concentrations of minerals can provide information about the capacity for uptake and translocation, it is well recognized that minerals are distributed heterogeneously across different cell types [7]. Not only may the total concentrations vary at the tissue, cellular, and subcellular scales but also the chemical speciation of minerals may vary. This spatial information is crucial for understanding the homeostasis of minerals, particularly how different cell types and, fundamentally, different genes function in controlling the distribution, complexation, and storage of minerals, and how these processes vary among diverse plant species in the ecophysiological context.A range of techniques are available for mapping element distribution at various spatial scales. Traditional methods, such as energy-dispersive X-ray microanalysis (EDX) and proton (particle)-induced X-ray emission (PIXE), have been u...