Abstract
Phosphorus is a crucial macronutrient for plants playing a critical role in many cellular signaling and energy cycling processes. In light of this, phosphorus acquisition efficiency is an important target trait for crop improvement, but it also provides an ecological adaptation for growth of plants in low nutrient environments. Increased root hair density has been shown to improve phosphorus uptake and plant health in a number of species. In several plant families, including Brassicaceae, root hair bearing cells are positioned on the epidermis according to their position in relation to cortex cells, with hair cells positioned in the cleft between two underlying cortex cells. Thus the number of cortex cells determines the number of epidermal cells in the root hair position. Previous research has associated phosphorus-limiting conditions with an increase in the number of cortex cell files in Arabidopsis thaliana roots, but they have not investigated the spatial or temporal domains in which these extra divisions occur or explored the consequences this has had on root hair formation. In this study, we use 3D reconstructions of root meristems to demonstrate that the radial anticlinal cell divisions seen under low phosphate are exclusive to the cortex. When grown on media containing replete levels of phosphorous, A. thaliana plants almost invariably show eight cortex cells; however when grown in phosphate limited conditions, seedlings develop up to 16 cortex cells (with 10–14 being the most typical). This results in a significant increase in the number of epidermal cells at hair forming positions. These radial anticlinal divisions occur within the initial cells and can be seen within 24 h of transfer of plants to low phosphorous conditions. We show that these changes in the underlying cortical cells feed into epidermal patterning by altering the regular spacing of root hairs.
Highlights
IntroductionUnlike nitrogen or ammonium, which are soluble and diffuse through the soil, phosphate (Pi) is immobile within the soil as it often binds clay particles and forms insoluble precipitates (Brady and Weil, 2008)
Phosphorus (P) is a critical macronutrient and essential for plant growth
Previous reports have shown that cortical cell proliferation is increased when A. thaliana plants are grown on low phosphate media (Ma et al, 2003; Cederholm and Benfey, 2015), it has been unclear whether the cortex is the only cell type to undergo additional radial anticlinal divisions to increase cell file number
Summary
Unlike nitrogen or ammonium, which are soluble and diffuse through the soil, phosphate (Pi) is immobile within the soil as it often binds clay particles and forms insoluble precipitates (Brady and Weil, 2008). Tomato, spinach, rape and Arabidopsis all exhibit an increase in root hair length under low Pi (Foehse and Jungk, 1983; Ma et al, 2001; Bhosale et al, 2018). These species respond by increasing their root hair density in low Pi conditions. A recent study of root hair traits in 166 accessions of Arabidopsis thaliana showed that root hair density and length were not correlated, and that the genotypes that showed the greatest increase in root hair density under low Pi were mostly those that had fewer and shorter root hairs under Pi replete conditions (Stetter et al, 2015)
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