doi: 10.1098/rstb.1993.0086, Clough, S. J., Fengler, K. A., Yu, I.-C., Lippok, B., Smith, R. K., and Bent, A. F. (2000). (2013). Plant Physiol. Interplay between ions, the cytoskeleton, and cell wall properties during tip growth. To exert this function, free Ca2+ levels in the cytosol under non-stimulated conditions need to be kept at a low level of around 0.1 μM. doi: 10.1093/jexbot/52.358.891, White, P. J., Bowen, H. C., Demidchik, V., Nichols, C., and Davies, J. M. (2002). doi: 10.1111/j.1469-8137.2008.02682.x, Michard, E., Lima, P. T., Borges, F., Silva, A. C., Portes, M. T., Carvalho, J. E., et al. (2013). *Correspondence: Kathrin Thor, kathrin.thor@tsl.ac.uk, Front. Calcium. Proc. doi: 10.1105/tpc.11.4.691. (2012). Hepler, P. K. (2005). Opin. 147, 1690–1698. It is needed in large amounts by all plants for the formation of cell walls and cell membranes, and it plays a vital role in soil structure, according to an article on HelenaChemical.com. To fulfill the structural role, calcium has to be available for the plant in sufficient amounts. Plant J. Proc. 28, 666–675. Plant Cell 17, 2142–2155. Calcium signals: the lead currency of plant information processing. Field application of calcium to reduce phytophthora stem rot of soybean, and calcium distribution in plants. Genes for calcium-permeable channels in the plasma membrane of plant root cells. The CBL-interacting protein kinase CIPK23 regulates HAK5-mediated high-affinity K+ uptake in Arabidopsis roots. doi: 10.1016/j.chom.2014.10.007, Monshausen, G. B., Messerli, M. A., and Gilroy, S. (2008). However, in recent years, it has emerged that the main function of calcium is to serve as a second messenger. Once loaded into the xylem, calcium is transported to the shoot, where it finally has to be unloaded into and distributed within the leaf cells (White and Broadley, 2003). Acad. The author confirms being the sole contributor of this work and has approved it for publication. Copyright © 2019 Thor. (2018). These elevations were also shown to be dependent on the coreceptor BAK1 with the corresponding receptor perceiving the herbivore-associated molecular pattern still awaiting identification (Vincent et al., 2017). (2014). Cytosolic calcium signals elicited by the pathogen-associated molecular pattern flg22 in stomatal guard cells are of an oscillatory nature. The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Plant Cell 4, 1123–1130. As calcium signals are involved in a diverse range of processes, it is assumed that the spatiotemporal pattern of the calcium rise, the so-called Ca2+ signature, determines the specificity of the corresponding response (McAinsh and Pittman, 2009; Dodd et al., 2010). Bot. Biophys. Proc. doi: 10.1126/science.1201101, Miedema, H., Bothwell, J. H. F., Brownlee, C., and Davies, J. M. (2001). Bifurcation of Arabidopsis NLR immune signaling via Ca2+-dependent protein kinases. Calcium was deemed an essential element for plant growth and development well over a hundred years ago and its importance has been heavily documented. doi: 10.1111/j.1469-8137.2008.02465.x, Monaghan, J., Matschi, S., Shorinola, O., Rovenich, H., Matei, A., Segonzac, C., et al. Due to the immobility of calcium in the soil and plant tissues, a continuous supply must be present for plants to access. Calcium: a central regulator of plant growth and development. In general, calcium deficiency as a result of low soil availability is not very common (White and Broadley, 2003). Nature 549, 91–95. Plants are exposed to a variety of pathogens, ranging from bacteria to fungi and oomycetes, and are also attacked by insects. The pathways of calcium movement to the xylem. While CaM, CMLs, CBLs, and CDPKs contain EF-hand Ca2+-binding motifs, CIPKs are activated by the interaction with CBLs (Kudla et al., 2018). Wang et al. 136, 2438–2442. Because of its limited movement in the plant, adding calcium during critical development stages such as fruiting can vastly improve fruit quality, quantity and overall plant health. (1991). The primary function of calcium in plant growth is to provide structural support to cell walls. Function of Calcium. (2002). We now know that cell wall-degrading enzymes and cell wall fragments can induce signaling pathways, in which Ca2+ serves as second messenger. Proc. In these situations, it is important to add calcium with large amounts of gypsum or lime depending on other factors of soil chemistry. The RPM1 plant disease resistance gene facilitates a rapid and sustained increase in cytosolic calcium that is necessary for the oxidative burst and hypersensitive cell death. 4 doi: 10.1146/annurev-arplant-070109-104628, Dodds, P. N., and Rathjen, J. P. (2010). The transport proteins responsible for the uptake of K+, AKT1 (Arabidopsis K+ transporter 1), and HAK5 (high-affinity K+ transporter 5) in Arabidopsis are both regulated via the same Ca2+ decoding complex consisting of CBL1/9 and CIPK23 (Xu et al., 2006; Ragel et al., 2015; Behera et al., 2017). Cell 138, 1184–1194. (2013). Also, despite few examples, surprisingly little is known about how the involvement of calcium in one plant aspect is affecting another and whether the same or different genes are involved in two processes, as, for example, those taking up Ca2+ as a nutrient in roots and those responsible for influx during signal generation. (2017). Mol. Cell 69, 953–964. Nature 428, 764–767. In the form of Calcium Pectate, Calcium holds the cell walls of plants together. doi: 10.1104/pp.108.123638, Monshausen, G. B., Miller, N. D., Murphy, A. S., and Gilroy, S. (2011). Two spatially and temporally distinct Ca2+ signals convey Arabidopsis thaliana responses to K+ deficiency. Proteinaceous and oligosaccharidic elicitors induce different calcium signatures in the nucleus of tobacco cells. The signal is terminated by transport of Ca2+ out of the cytosol via Ca2+-ATPases (shown in orange) or H+/Ca2+ antiporters (purple) in the plasma or organeller membranes. 50, 1267–1280. McAinsh, M. R., and Pittman, J. K. (2009). CPK5 has recently been assigned a unique role in plant immunity in that, in contrast to CPK4, 6, or 11, it contributes to exo70B1-activated autoimmune responses via association with the truncated NLR protein TIR-NBS2 (TN2) (Liu et al., 2017). Plant Physiol. (2004). FERONIA receptor-like kinase regulates RHO GTpase signaling of root hair development. Schreiber, L., Hartmann, K., Skrabs, M., and Zeier, J. Yu, I.-c., Parker, J., and Bent, A. F. (1998). In Arabidopsis, they are recognized by the receptor Wall Associated Kinase 1 (WAK1) (He et al., 1996; Brutus et al., 2010). doi: 10.1111/tpj.12311. Influx into leaf cells after unloading from the vasculature in Arabidopsis has been proposed to occur via CNGC2. In today’s complex and fast-paced crop production sector, the team at CROPLIFE keeps 21,000 agricultural retailers, distributors and their suppliers up to date on such decidedly 21st century issues as seed technology, biotechnology, precision agriculture, customer service and retention, and business management. dnd1 mutants exhibit an autoimmune phenotype, being dwarf and showing constitutively elevated levels of salicylic acid (SA) as well as defense gene expression such as PR1 (Yu et al., 1998; Clough et al., 2000). In accordance with this, early studies reported a correlation of a low plant calcium tissue content with a higher fungal infection rate, suggesting that due to the lower membrane stability, the efflux of compounds such as sugars into the apoplast could be increased (Marschner, 1995). doi: 10.1038/nrg2812, Duan, Q., Kita, D., Li, C., Cheung, A. Y., and Wu, H.-M. (2010). In addition, calcium was reported to inhibit cell wall-degrading enzymes produced by fungi and bacteria, thus a lower calcium content could lead to higher degradation and infection (Marschner, 1995). (2006). 23, 441–450. Plant Physiol. This is because calcium displaces sodium in the soil, and with adequate leaching irrigations, it can help improve overall soil quality. doi: 10.1105/tpc.16.00822, Ma, Y., Walker, R. K., Zhao, Y., and Berkowitz, G. A. Figure 1. To protect themselves against this biotic stress, they have evolved a two-layered immune signaling network providing defense. (2015). Analysis and effects of cytosolic free calcium increases in response to elicitors in Nicotiana plumbaginifolia cells. doi: 10.1093/emboj/20.20.5556, Sanders, D., Brownlee, C., and Harper, J. F. (1999). 204, 873–881. Annu. Young developing tissues such as growing points and fruiting bodies tend to be affected the most once soil reserves are depleted.

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