Auxin is essential for the regulation of root system architecture by controlling primary root elongation and lateral root (LR) formation. Exogenous auxin has been reported to inhibit primary root elongation and promote the formation of LRs. In this study, LR formation in the Zea mays primary root was quantitatively evaluated after exogenous auxin treatment by comparing the effects of auxin on two selected zones elongated either before or after auxin application. We determined two main variables in both zones: the LR density per unit of root length (LRD), and the mean phloem pericycle cell length. The total number of phloem pericycle cells (PPCs) per unit of root length was then calculated. Considering that each LR primordium is initiated from four founder cells (FCs), the percentage of PPCs (%PPC) that behave as FCs in a specific root zone was estimated by dividing the number of pericycle cells by four times the LRD. This index was utilized to describe LR initiation. Root zones elongated in the presence of a synthetic auxin (1-naphthalene acetic acid, NAA) at low concentrations (0.01 μM) showed reduced cell length and increased LRD. However, a high concentration of NAA (0.1 μM) strongly reduced both cell length and LRD. In contrast, both low and high levels of NAA stimulated LRD in zones elongated before auxin application. Analysis of the percentage of FCs in the phloem pericycle in zones elongated in the presence or absence of NAA showed that low concentrations of NAA increased the %PFC, indicating that LR initiation is promoted at new sites; however, high concentrations of NAA elicited a considerable reduction in this variable in zones developed in the presence of auxin. As these zones are composed of short pericycle cells, we propose that short pericycle cells are incapable to participate in LR primordium initiation and that auxin modulated initiation of LRs is linked to pericycle cell length.

The root system is responsible for water and nutrients uptake from the soil, and therefore, its extension is basic for an efficient acquisition. The maize root system is formed by different types of roots, and the lateral root branching substantially increases the surface for nutrient uptake. Therefore, the regulation of lateral root formation is fundamental in the development of root functions. Root architecture is basically controlled by auxin and cytokinins, which antagonize in the formation of lateral roots (LR) along the primary root axis, with auxin, a stimulator, and cytokinins inhibitors of LR development. This interaction has been analyzed in several zones along the primary root where LRs in different developmental stages were located. The root has been divided into several zones, such as meristem, elongation zone, and mature zone, according to the developmental processes occurring in each one. As Arabidopsis root elongated more slowly than maize root, these zones are shorter, and its delimitation is more difficult. However, these zones have previously been delimitated clearly in maize, and therefore, they analyze the effect of exogenous hormones in several LR developmental stages. The inhibitory effect of cytokinin on lateral root formation was observed in already elongated primary root zones in which initial events to form new lateral roots are taking place. Contrarily, auxin increased LR formation in the primary root segments elongated in the presence of the hormone. The inhibitory effect of cytokinin was reversed by auxin in a concentration-dependent manner when both hormones were combined. However, auxin is unable to recover LR development in primary root zones that have been previously elongated only in the presence of cytokinin. This antagonistic auxin-cytokinin effect on LR development depended on the balance between both hormones, which controls the root system architecture and determines the formation of LR during the process of initiation.

During the first days of development, maize roots showed considerable variation in the production of ethylene and the rate of elongation. As endogenous ethylene increases, root elongation decreases...

The role of auxins on root system architecture was studied by applying indole-3-acetic acid (IAA), indole-3-butyric acid (IBA), and 1-naphthaleneacetic acid (NAA) to maize roots and analysing the main processes involved in root development: primary root (PR) elongation, lateral root (LR) formation, and LR root elongation. We found that these effects were not dependent only on concentration, but also on the type of auxin applied. We also studied temporal changes in auxin inhibition of PR elongation. These temporal changes were analysed calculating the elongation ratio between two consecutive one day periods after auxin application. It was observed that a reduction in root elongation was also dependent on the type of auxin applied and its concentration. The inhibitory effect of IBA and IAA decreased on the second day, and the ratio also increased with the concentration. In contrast, NAA increased root elongation inhibition with time. Indeed, the ratio decreased as the NAA concentration increased. Regarding LR formation, we observed that external auxin increased only LR formation in certain zones of the PR. Finally, comparison of inhibition elongation associated with auxin in the LR and PR clearly demonstrates that PR elongation was more sensitive to auxin than LR elongation.

Auxin and ethylene have been largely reported to reduce root elongation in maize primary root. However the effects of auxin are greater than those caused by ethylene. Although auxin stimulates ethylene biosynthesis through the specific increase of ACC synthase, the auxin inhibitory effect on root elongation is not mediated by the auxin-induced increase of ethylene production. Recently it has been demonstrated that root inhibition by the application of the synthetic auxin NAA (1-naphtalenacetic acid) is increased if combined with the ethylene precursor ACC (1-aminocyclopropane-1-carboxilic acid) when both compounds are applied at very low concentrations.