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The inhibitory effect of ammonium on primary root growth has been well documented; however the underlying physiological and molecular mechanisms are still controversial. To avoid ammonium toxicity to shoot growth, we used a vertical two-layer split plate system, in which the upper layer contained nitrate and the lower layer contained ammonium. In this way, nitrogen status was maintained and only the apical part of the root system was exposed to ammonium. Using a kinematic approach, we show here that 1 mM ammonium reduces primary root growth, decreasing both elemental expansion and cell production. Ammonium inhibits the length of elongation zone and the maximum elemental expansion rate. Ammonium also decreases the apparent length of the meristem as well as the number of dividing cells without affecting cell division rate. Moreover, ammonium reduces the number of root cap cells but appears to affect neither the status of root stem cell niche nor the distal auxin maximum at the quiescent center. Ammonium also inhibits root gravitropism and concomitantly down-regulates the expression of two pivotal auxin transporters, AUX1 and PIN2. Insofar as ammonium inhibits root growth rate in AUX1 and PIN2 loss-of-function mutants almost as strongly as in wild type, we conclude that ammonium inhibits root growth and gravitropism by largely distinct pathways.

HighlightThis research determined the significant genetic and phenotypic relationships between seedling root traits and nitrogen use efficiency (NUE), and further identified five QTL clusters for improving NUE in maize.

Under low-input cropping systems, nitrogen (N) can be a limiting factor in plant growth and yield. Identifying genotypes that are more efficient at capturing limited N resources and the traits and mechanisms responsible for this ability is important. Root trait has a substantial influence on N acquisition from soils. Nevertheless, inconsistencies still exist as to the effect of low N on root length and its architecture in terms of lateral and axial roots. For maize, a crop utilizing heterosis, little is known about the relationship between parents and their crosses in the response of root architecture to N availability. Here 7 inbred maize lines and 21 of their crosses created by diallel mating were used to study the effect of N stress on root morphology as well as the relationship between the inbreds and their crosses. With large genotypic differences, low N generally suppresses shoot growth and increases the root to shoot ratio with or without increasing root biomass in maize. Maize plants responded to N deficiency by increasing total root length and altering root architecture by increasing the elongation of individual axial roots and enhancing lateral root growth, but with a reduction in the number of axial roots. Here, the inbreds showed weaker responses in root biomass and other root parameters than their crosses. Heterosis of root traits was significant at both N levels and was attributed to both the general combining ability (GCA) and special combining ability (SCA). Low N had substantial affects on the pattern of heterosis, GCA and SCA affects on root traits for each of the crosses suggesting that selection under N stress is necessary in generating low N-tolerant maize genotypes.

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The plasticity of root architecture is crucial for plants to acclimate to unfavourable environments including low nitrogen (LN) stress. How maize roots coordinate the growth of axile roots and lateral roots (LRs), as well as longitudinal and radial cell behaviours in response to LN stress, remains unclear. Maize plants were cultivated hydroponically under control (4 mm nitrate) and LN (40 μm) conditions. Temporal and spatial samples were taken to analyse changes in the morphology, anatomical structure and carbon/nitrogen (C/N) ratio in the axile root and LRs. LN stress increased axile root elongation, reduced the number of crown roots and decreased LR density and length. LN stress extended cell elongation zones and increased the mature cell length in the roots. LN stress reduced the cell diameter and total area of vessels and increased the amount of aerenchyma, but the number of cell layers in the crown root cortex was unchanged. The C/N ratio was higher in the axile roots than in the LRs. Maize roots acclimate to LN stress by optimizing the anatomical structure and N allocation. As a result, axile root elongation is favoured to efficiently find available N in the soil.

Nitrogen (N) loss is a worldwide problem in crop production. Apart from reasonable N fertilizer application, breeding N efficient cultivars provides an alternative way. Root architecture is an important factor determining N acquisition. However, little is known about the molecular genetic basis for root growth in relation to N supply. In the present study, an F 8 maize (Zea may L.) recombinant inbred (RI) population consisting of 94 lines was used to identify the QTLs for root traits under different nitrate levels. The lateral root length (LRL), axial root length (ARL), maximal axial root length (MARL), axial root number (ARN) and average axial root length (AARL) were evaluated under low N (LN) and high N (HN) conditions in a hydroponics system. A total of 17 QTLs were detected among which 14 loci are located on the same chromosome region as published QTLs for root traits. A major QTL on chromosome 1 (between bnlg1025 and umc2029) for the AARL under LN could explain 43.7% of the phenotypic variation. This QTL co-localizes with previously reported QTLs that associate with root traits, grain yield, and N uptake. Our results indicate that longer axial roots are important for efficient N acquisition and the major QTL for AARL may be used as a marker in breeding N efficient maize genotypes.