Cortex cell hydraulic conductivity, endodermal apoplastic barriers and root hydraulics change in barley (Hordeum vulgare L.) in response to a low supply of N and P

Abstract

Background Mineral nutrient limitation affects the water flow through plants. We wanted to test on barley whether any change in root-to-shoot ratio in response to low supply of nitrogen and phosphate is accompanied by changes in root and cell hydraulic properties and involves changes in aquaporin (AQP) gene expression and root apoplastic barriers (suberin lamellae, Casparian bands). Methods Plants were grown hydroponically on complete nutrient solution or on solution containing only 3.3 % or 2.5 % of the control level of nutrient. Plants were analysed when they were 14–18 d old. Results Nutrient-limited plants adjusted water flow to an increased root-to-shoot surface area ratio through a reduction in root hydraulic conductivity (Lp) as determined through exudation analyses. Cortex cell Lp (cell pressure probe analyses) decreased in the immature but not the mature region of the main axis of seminal roots and in primary lateral roots. The aquaporin inhibitor HgCl2 reduced root Lp most in nutrient-sufficient control plants. Exchange of low-nutrient for control media caused a rapid (20–80 min) and partial recovery in Lp, though cortex cell Lp did not increase in any of the root regions analysed. The gene expression level (qPCR analyses) of five plasma membrane-localized AQP isoforms did not change in bulk root extracts, while the formation of apoplastic barriers increased considerably along the main axis of root and lateral roots in low-nutrient treatments. Conclusions Decrease in root and cortex cell Lp enables the adjustment of root water uptake to increased root-to-shoot area ratio in nutrient-limited plants. Aquaporins are the prime candidate to play a key role in this response. Modelling of water flow suggests that some of the reduction in root Lp is due to increased formation of apoplastic barriers.

Exodermis and Endodermis Respond to Nutrient Deficiency in Nutrient-Specific and Localized Manner

“…Its suberization was enhanced in -N or -P roots but delayed in -K roots. These trends generally correspond with previously documented changes in overall radial water flow in wheat, sunflower, and maize; decreased radial water transport under N, P, or S deficiency and increased transport under K deficiency [31,[38][39][40]. However, there are also studies showing the opposite trends, e.g., decreased root hydraulic conductivity in low-K barley [41].…”

“…We therefore extended our examination to barley with the same result of enhanced exodermal differentiation in -P compared to control conditions. Recently, this response of barley was presented in another study [31].…”

“…This can be a way to regulate water transport [31,48]. Low hydraulic conductivity of deficient roots might adjust water uptake to higher root/shoot ratio of deficient plants [31]. However, we rather propose that this response might be the preference to prevent nutrient leakage from the stele and cortex back to the rhizosphere over leaving the apoplastic path free for nutrient uptake from the rhizosphere.…”

“…All these results taken together show that the examined grass crop species respond to severe N or P limitation (zero N or P; strong signs of deficiency on aerial parts) by enhanced differentiation of apoplastic barriers, both in the exodermis and endodermis, in our study. This can be a way to regulate water transport [31,48]. Low hydraulic conductivity of deficient roots might adjust water uptake to higher root/shoot ratio of deficient plants [31].…”

“…Three wetland Carex species (C. vesicaria, C. rostrata, and C. gracilis) showed enhanced exodermis differentiation in oligotrophic compared to eutrophic conditions in sand culture [29]; enhanced exodermal suberization was found in maize under Mg deficiency [30]. Armand (2019) recently emphasised that enhanced endodermal and exodermal differentiation in low-N and low-P barley corresponds with reduced root hydraulic conductivity [31].…”