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Plants generally enhance their root growth in the form of greater biomass and/or root length to boost nutrient uptake in response to short-term low nitrogen (LN). However, the underlying mechanisms of short-term LN-mediated root growth remain largely elusive. Our genome-wide association study, haplotype analysis, and phenotyping of transgenic plants showed that the crucial nitrate signaling component NIN-LIKE PROTEIN3.2 (ZmNLP3.2), a positive regulator of root biomass, is associated with natural variations in root biomass of maize (Zea mays L.) seedlings under LN. The monocot-specific gene AUXIN/INDOLE-3-ACETIC ACID14 (ZmAux/IAA14) exhibited opposite expression patterns to ZmNLP3.2 in ZmNLP3.2 knockout and overexpression lines, suggesting that ZmNLP3.2 hampers ZmAux/IAA14 transcription. Importantly, ZmAux/IAA14 knockout seedlings showed a greater root dry weight (RDW), whereas ZmAux/IAA14 overexpression reduced RDW under LN compared with wild-type plants, indicating that ZmAux/IAA14 negatively regulates the RDW of LN-grown seedlings. Moreover, in vitro and vivo assays indicated that AUXIN RESPONSE FACTOR19 (ZmARF19) binds to and transcriptionally activates ZmAux/IAA14, which was weakened by the ZmNLP3.2-ZmARF19 interaction. The zmnlp3.2 ZmAux/IAA14-OE seedlings exhibited further reduced RDW compared with ZmAux/IAA14 overexpression lines when subjected to LN treatment, corroborating the ZmNLP3.2-ZmAux/IAA14 interaction. Thus, our study reveals a ZmNLP3.2-ZmARF19-ZmAux/IAA14 module regulating root biomass in response to nitrogen limitation in maize., Competing Interests: Conflict of interest statement. The authors declare no conflict of interests., (© The Author(s) 2024. Published by Oxford University Press on behalf of American Society of Plant Biologists. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Plant phenotypic plasticity plays an important role in nitrogen (N) acquisition and use under nitrogen-limited conditions. However, this role has never been quantified as a function of N availability, leaving it unclear whether plastic responses should be considered as potential targets for selection. A combined modelling and experimentation approach was adopted to quantify the role of plasticity in N uptake and plant yield. Based on a greenhouse experiment we considered plasticity in two maize (Zea mays) traits: root-to-leaf biomass allocation ratio and emergence rate of axial roots. In a simulation experiment we individually enabled or disabled both plastic responses for maize stands grown across six N levels. Both plastic responses contributed to maintaining a higher N uptake, and plant productivity as N availability declined compared with stands in which plastic responses were disabled. We conclude that plastic responses quantified in this study may be a potential target trait in breeding programs for greater N uptake across N levels while it may only be important for the internal use of N under N-limited conditions in maize. Given the complexity of breeding for plastic responses, an a priori model analysis is useful to identify which plastic traits to target for enhanced plant performance., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Under low nitrogen (N) supply, an important adaption of the maize root system is to promote the root elongation so as to increase N uptake from a larger soil space. The underlying physiological mechanism is largely unknown. In the present study, two maize inbred lines (Ye478 and Wu312) were used to study the possible involvement of the auxin and target of rapamycin (TOR) pathway in low-N-induced root elongation. Compared to Wu312, primary root elongation of Ye478 was more sensitive to low nitrate supply. Correspondingly, more auxin was accumulated in the root tip, and more protons were secreted, increasing the acidity of the apoplast space. On the other hand, low-N-induced root elongation was greatly reduced when shoot-to-root auxin transport was inhibited by applying N-1-naphthylphthalamic acid (NPA) at the plant base or by pruning the top leaf where auxin is mostly synthesized. Furthermore, exogenous application of TOR inhibitor also eliminated the response of root elongation under low N. The content of TOR kinase and the expression of TOR pathway-related genes were significantly changed when shoot-to-root auxin transport was reduced by NPA treatment. Taken together, it is concluded that low-N stress increases shoot-to-root auxin transport which enhances root elongation via auxin-dependent acid growth and the auxin-regulated TOR pathway in maize., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

Main Conclusion: In response to low nitrogen stress, multiple hormones together with nitric oxide signaling pathways work synergistically and antagonistically in crop root elongation. Changing root morphology allows plants to adapt to soil nutrient availability. Nitrogen is the most important essential nutrient for plant growth. An important adaptive strategy for crops responding to nitrogen deficiency is root elongation, thereby accessing increased soil space and nitrogen resources. Multiple signaling pathways are involved in this regulatory network, working together to fine-tune root elongation in response to soil nitrogen availability. Based on existing research, we propose a model to explain how different signaling pathways interact to regulate root elongation in response to low nitrogen stress. In response to a low shoot nitrogen status signal, auxin transport from the shoot to the root increases. High auxin levels in the root tip stimulate the production of nitric oxide, which promotes the synthesis of strigolactones to accelerate cell division. In this process, cytokinin, ethylene, and abscisic acid play an antagonistic role, while brassinosteroids and auxin play a synergistic role in regulating root elongation. Further study is required to identify the QTLs, genes, and favorable alleles which control the root elongation response to low nitrogen stress in crops.

Ammonium uptake in plant roots is mediated by AMT/MEP/Rh-type ammonium transporters. Out of five AMTs being expressed in Arabidopsis roots, four AMT1-type transporters contribute to ammonium uptake, whereas no physiological function has so far been assigned to the only homolog belonging to the MEP subfamily, AMT2;1. Based on the observation that under ammonium supply, the transcript levels of AMT2;1 increased and its promoter activity shifted preferentially to the pericycle, we assessed the contribution of AMT2;1 to xylem loading. When exposed to 15 N-labeled ammonium, amt2;1 mutant lines translocated less tracer to the shoots and contained less ammonium in the xylem sap. Moreover, in an amt1;1 amt1;2 amt1;3 amt2;1 quadruple mutant (qko), co-expression of AMT2;1 with either AMT1;2 or AMT1;3 significantly enhanced 15 N translocation to shoots, indicating a cooperative action between AMT2;1 and AMT1 transporters. Under N deficiency, proAMT2;1-GFP lines showed enhanced promoter activity predominantly in cortical root cells, which coincided with elevated ammonium influx conferred by AMT2;1 at millimolar substrate concentrations. Our results indicate that in addition to contributing moderately to root uptake in the low-affinity range, AMT2;1 functions mainly in root-to-shoot translocation of ammonium, depending on its cell-type-specific expression in response to the plant nutritional status and to local ammonium gradients., (Copyright © 2017 The Authors. Published by Elsevier Inc. All rights reserved.)

Root system architecture (RSA) plays an important role in phosphorus (P) acquisition, but enhancing P use efficiency (PUE) in maize via genetic manipulation of RSA has not yet been reported. Here, using a maize recombinant inbred line (RIL) population, we investigated the genetic relationships between PUE and RSA, and developed P-efficient lines by selection of quantitative trait loci (QTLs) that coincide for both traits. In low-P (LP) fields, P uptake efficiency (PupE) was more closely correlated with PUE (r = 0.48-0.54), and RSA in hydroponics was significantly related to PupE (r = 0.25-0.30) but not to P utilization efficiency (PutE). QTL analysis detected a chromosome region where two QTLs for PUE, three for PupE and three for RSA were assigned into two QTL clusters, Cl-bin3.04a and Cl-bin3.04b. These QTLs had favorable effects from alleles derived from the large-rooted and high-PupE parent. Marker-assisted selection (MAS) identified nine advanced backcross-derived lines carrying Cl-bin3.04a or Cl-bin3.04b that displayed mean increases of 22%-26% in PUE in LP fields. Furthermore, a line L224 pyramiding Cl-bin3.04a and Cl-bin3.04b showed enhanced PupE, relying mainly on changes in root morphology, rather than root physiology, under both hydroponic and field conditions. These results highlight the physiological and genetic contributions of RSA to maize PupE, and provide a successful study case of developing P-efficient crops through QTL-based selection., (Copyright © 2016 Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, and Genetics Society of China. Published by Elsevier Ltd. All rights reserved.)

A plant's ability to maintain or improve its yield under limiting conditions, such as nutrient deficiency or drought, can be strongly influenced by root system architecture (RSA), the three-dimensional distribution of the different root types in the soil. The ability to image, track and quantify these root system attributes in a dynamic fashion is a useful tool in assessing desirable genetic and physiological root traits. Recent advances in imaging technology and phenotyping software have resulted in substantive progress in describing and quantifying RSA. We have designed a hydroponic growth system which retains the three-dimensional RSA of the plant root system, while allowing for aeration, solution replenishment and the imposition of nutrient treatments, as well as high-quality imaging of the root system. The simplicity and flexibility of the system allows for modifications tailored to the RSA of different crop species and improved throughput. This paper details the recent improvements and innovations in our root growth and imaging system which allows for greater image sensitivity (detection of fine roots and other root details), higher efficiency, and a broad array of growing conditions for plants that more closely mimic those found under field conditions., (© 2015 Institute of Botany, Chinese Academy of Sciences.)

Maize (Zea mays L.) root morphology exhibits a high degree of phenotypic plasticity to nitrogen (N) deficiency, but the underlying genetic architecture remains to be investigated. Using an advanced BC4 F3 population, we investigated the root growth plasticity under two contrasted N levels and identified the quantitative trait loci (QTLs) with QTL-environment (Q × E) interaction effects. Principal components analysis (PCA) on changes of root traits to N deficiency (ΔLN-HN) showed that root length and biomass contributed for 45.8% in the same magnitude and direction on the first PC, while root traits scattered highly on PC2 and PC3. Hierarchical cluster analysis on traits for ΔLN-HN further assigned the BC4 F3 lines into six groups, in which the special phenotypic responses to N deficiency was presented. These results revealed the complicated root plasticity of maize in response to N deficiency that can be caused by genotype-environment (G × E) interactions. Furthermore, QTL mapping using a multi-environment analysis identified 35 QTLs for root traits. Nine of these QTLs exhibited significant Q × E interaction effects. Taken together, our findings contribute to understanding the phenotypic and genotypic pattern of root plasticity to N deficiency, which will be useful for developing maize tolerance cultivars to N deficiency., (© 2015 Institute of Botany, Chinese Academy of Sciences.)

That root system architecture (RSA) has an essential role in nitrogen acquisition is expected in maize, but the genetic relationship between RSA and nitrogen use efficiency (NUE) traits remains to be elucidated. Here, the genetic basis of RSA and NUE traits was investigated in maize using a recombination inbred line population that was derived from two lines contrasted for both traits. Under high-nitrogen and low-nitrogen conditions, 10 NUE- and 9 RSA-related traits were evaluated in four field environments and three hydroponic experiments, respectively. In contrast to nitrogen utilization efficiency (NutE), nitrogen uptake efficiency (NupE) had significant phenotypic correlations with RSA, particularly the traits of seminal roots (r = 0.15-0.31) and crown roots (r = 0.15-0.18). A total of 331 quantitative trait loci (QTLs) were detected, including 184 and 147 QTLs for NUE- and RSA-related traits, respectively. These QTLs were assigned into 64 distinct QTL clusters, and ~70% of QTLs for nitrogen-efficiency (NUE, NupE, and NutE) coincided in clusters with those for RSA. Five important QTLs clusters at the chromosomal regions bin1.04, 2.04, 3.04, 3.05/3.06, and 6.07/6.08 were found in which QTLs for both traits had favourable effects from alleles coming from the large-rooted and high-NupE parent. Introgression of these QTL clusters in the advanced backcross-derived lines conferred mean increases in grain yield of ~14.8% for the line per se and ~15.9% in the testcross. These results reveal a significant genetic relationship between RSA and NUE traits, and uncover the most promising genomic regions for marker-assisted selection of RSA to improve NUE in maize., (© The Author 2015. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

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., (© 2014 John Wiley & Sons Ltd.)

Approximately 35-55% of total nitrogen (N) in maize plants is taken up by the root at the reproductive stage. Little is known about how the root of an adult plant responds to heterogeneous nutrient supply. In this study, root morphological and physiological adaptations to nitrate-rich and nitrate-poor patches and corresponding gene expression of ZmNrt2.1 and ZmNrt2.2 of maize seedlings and adult plants were characterized. Local high nitrate (LoHN) supply increased both lateral root length (LRL) and density of the treated nodal roots of adult maize plants, but only increased LRL of the treated primary roots of seedlings. LoHN also increased plant total N acquisition but not N influx rate of the treated roots, when expressed as per unit of root length. Furthermore, LoHN markedly increased specific root length (m g(-1)) of the treated roots but significantly inhibited the growth of the lateral roots outside of the nitrate-rich patches, suggesting a systemic carbon saving strategy within a whole root system. Surprisingly, local low nitrate (LoLN) supply stimulated nodal root growth of adult plants although LoLN inhibited growth of primary roots of seedlings. LoLN inhibited the N influx rate of the treated roots and did not change plant total N content. The gene expression of ZmNrt2.1 and ZmNrt2.2 of the treated roots of seedlings and adult plants was inhibited by LoHN but enhanced by LoLN. In conclusion, maize adult roots responded to nitrate-rich and nitrate-poor patches by adaptive morphological alterations and displayed carbon saving strategies in response to heterogeneous nitrate supply., (© 2013 Scandinavian Plant Physiology Society.)

High-affinity ammonium uptake in plant roots is mainly mediated by AMT1-type ammonium transporters, and their regulation varies depending on the plant species. In this study we aimed at characterizing AMT-mediated ammonium transport in maize, for which ammonium-based fertilizer is an important nitrogen (N) source. Two ammonium transporter genes, ZmAMT1;1a and ZmAMT1;3, were isolated from a maize root-specific cDNA library by functional complementation of an ammonium uptake-defective yeast mutant. Ectopic expression of both genes in an ammonium uptake-defective Arabidopsis mutant conferred high-affinity ammonium uptake capacities in roots with substrate affinities of 48 and 33 μM for ZmAMT1;1a and ZmAMT1;3, respectively. In situ hybridization revealed co-localization of both ZmAMT genes on the rhizodermis, suggesting an involvement in capturing ammonium from the rhizosphere. In N-deficient maize roots, influx increased significantly while ZmAMT expression did not. Ammonium resupply to N-deficient or nitrate-pre-cultured roots, however, rapidly enhanced both influx and ZmAMT transcript levels, revealing a substrate-inducible regulation of ammonium uptake. In conclusion, the two rhizodermis-localized transporters ZmAMT1;1a and ZmAMT1;3 are most probably the major components in the high-affinity transport system in maize roots. A particular regulatory feature is their persistent induction by ammonium rather than an up-regulation under N deficiency.

Root and rhizosphere research has been conducted for many decades, but the underlying strategy of root/rhizosphere processes and management in intensive cropping systems remain largely to be determined. Improved grain production to meet the food demand of an increasing population has been highly dependent on chemical fertilizer input based on the traditionally assumed notion of 'high input, high output', which results in overuse of fertilizers but ignores the biological potential of roots or rhizosphere for efficient mobilization and acquisition of soil nutrients. Root exploration in soil nutrient resources and root-induced rhizosphere processes plays an important role in controlling nutrient transformation, efficient nutrient acquisition and use, and thus crop productivity. The efficiency of root/rhizosphere in terms of improved nutrient mobilization, acquisition, and use can be fully exploited by: (1) manipulating root growth (i.e. root development and size, root system architecture, and distribution); (2) regulating rhizosphere processes (i.e. rhizosphere acidification, organic anion and acid phosphatase exudation, localized application of nutrients, rhizosphere interactions, and use of efficient crop genotypes); and (3) optimizing root zone management to synchronize root growth and soil nutrient supply with demand of nutrients in cropping systems. Experiments have shown that root/rhizosphere management is an effective approach to increase both nutrient use efficiency and crop productivity for sustainable crop production. The objectives of this paper are to summarize the principles of root/rhizosphere management and provide an overview of some successful case studies on how to exploit the biological potential of root system and rhizosphere processes to improve crop productivity and nutrient use efficiency.

Interspecific root/rhizosphere interactions affect phosphorus (P) uptake and the productivity of maize/faba bean and maize/wheat intercropping systems. The aim of these experiments was to determine whether manipulation of maize root growth could improve the productivity of the two intercropping systems. Two near isogenic maize hybrids (the larger-rooted T149 and smaller-rooted T222) were intercropped with faba bean and wheat, under conditions of high- and low-P availability. The larger-rooted T149 showed greater competitive ability than the smaller-rooted T222 in both maize/faba bean and maize/wheat intercropping systems. The higher competitive ability of T149 improved the productivity of the maize/faba bean intercropping system in P-sufficient conditions. In maize/wheat intercropping systems, root growth, shoot biomass, and P uptake of maize were inhibited by wheat, regardless of the P-supply. Compared with T222, the larger-rooted T149 suffered less in the intercropping systems. The total biomass of the maize/wheat intercropping system was higher for wheat/T149 than for wheat/T222 under low-P conditions. These data suggested that genetic improvement of maize root size could enhance maize growth and its ability to compete for P resources in maize/faba bean and maize/wheat intercropping systems. In addition, depending on the P availability, larger maize roots could increase the productivity of intercropping systems.

Root system architecture (RSA) is seldom considered as a selection criterion to improve yield in maize breeding, mainly because of the practical difficulties with their evaluation under field conditions. In the present study, phenotypic profiling of 187 advanced-backcross BC(4)F(3) maize lines (Ye478 × Wu312) was conducted at different developmental stages under field conditions at two locations (Dongbeiwang in 2007 and Shangzhuang in 2008) for five quantitative root traits. The aims were to (1) understand the genetic basis of root growth in the field; (2) investigate the contribution of root traits to grain yield (GY); and (3) detect QTLs controlling root traits at the seedling (I), silking (II) and maturation (III) stages. Axial root (AR)-related traits showed higher heritability than lateral root (LR)-related traits, which indicated stronger environmental effects on LR growth. Among the three developmental stages, root establishment at stage I showed the closest relationship with GY (r = 0.33-0.43, P < 0.001). Thirty QTLs for RSA were detected in the BC(4)F(3) population and only 13.3 % of the QTLs were detected at stage III. Most important QTLs for root traits were located on chromosome 6 near the locus umc1257 (bin 6.02-6.04) at stage I, and chromosome 10 near the locus umc2003 (bin 10.04) for number of AR across all three developmental stages. The regions of chromosome 7 near the locus bnlg339 (bin 7.03) and chromosome 1 near the locus bnlg1556 (bin 1.07) harbored QTLs for both GY- and LR-related traits at stages I and II, respectively. These results help to understand the genetic basis of root development under field conditions and their contribution to grain yield.

Root growth has a fundamental role in nitrogen (N) use efficiency. Nevertheless, little is known about how modern breeding progress has affected root growth and its responses to N supply. The root and shoot growth of a core set of 11 representative Chinese maize (Zea mays L.) hybrids released between 1973 and 2009 were investigated under high N (4 mmol L(-1), HN) and low N (0.04 mmol L(-1), LN) levels in a solution culture system. Compared with LN, HN treatment decreased root dry weight (RDW), the root: shoot ratio (R/S), and the relative growth rate for root dry weight (RGR(root)), but increased the total root length (TRL) and the total lateral root length (LRL). The total axial root length (ARL) per plant was reduced under HN, mostly in hybrids released before the 1990s. The number of seminal roots (SRN) was largely unaffected by different N levels. More recently released hybrids showed higher relative growth rates in the shoot under both HN and LN. However, the roots only showed increased RGR under HN treatment. Correspondingly, there was a positive linear relationship with the year of hybrid release for TRL, LRL and ARL under HN treatment. Together, these results suggest that while shoot growth of maize has improved, its root growth has only improved under high N conditions over the last 36 years of selective breeding in China. Improving root growth under LN conditions may be necessary to increase the N use efficiency of maize.

Unlabelled: background and aims: Roots typically respond to localized nitrate by enhancing lateral-root growth. Polar auxin transport has important roles in lateral-root formation and growth; however, it is a matter of debate whether or how auxin plays a role in the localized response of lateral roots to nitrate., Methods: Treating maize (Zea mays) in a split-root system, auxin levels were quantified directly and polar transport was assayed by the movement of [(3)H]IAA. The effects of exogenous auxin and polar auxin transport inhibitors were also examined., Key Results: Auxin levels in roots decreased more in the nitrate-fed compartment than in the nitrate-free compartment and nitrate treatment appeared to inhibit shoot-to-root auxin transport. However, exogenous application of IAA only partially reduced the stimulatory effect of localized nitrate, and auxin level in the roots was similarly reduced by local applications of ammonium that did not stimulate lateral-root growth., Conclusions: It is concluded that local applications of nitrate reduced shoot-to-root auxin transport and decreased auxin concentration in roots to a level more suitable for lateral-root growth. However, alteration of root auxin level alone is not sufficient to stimulate lateral-root growth.

The use of nitrogen (N) fertilizers has contributed to the production of a food supply sufficient for both animals and humans despite some negative environmental impact. Sustaining food production by increasing N use efficiency in intensive cropping systems has become a major concern for scientists, environmental groups, and agricultural policymakers worldwide. In high-yielding maize systems the major method of N loss is nitrate leaching. In this review paper, the characteristic of nitrate movement in the soil, N uptake by maize as well as the regulation of root growth by soil N availability are discussed. We suggest that an ideotype root architecture for efficient N acquisition in maize should include (i) deeper roots with high activity that are able to uptake nitrate before it moves downward into deep soil; (ii) vigorous lateral root growth under high N input conditions so as to increase spatial N availability in the soil; and (iii) strong response of lateral root growth to localized nitrogen supply so as to utilize unevenly distributed nitrate especially under limited N conditions.

Despite the fact that urea is a ubiquitous nitrogen source in soils and the most widespread form of nitrogen fertilizer used in agricultural plant production, membrane transporters that might contribute to the uptake of urea in plant roots have so far been characterized only in heterologous systems. Two T-DNA insertion lines, atdur3-1 and atdur3-3, that showed impaired growth on urea as a sole nitrogen source were used to investigate a role of the H+/urea co-transporter AtDUR3 in nitrogen nutrition in Arabidopsis. In transgenic lines expressing AtDUR3-promoter:GFP constructs, promoter activity was upregulated under nitrogen deficiency and localized to the rhizodermis, including root hairs, as well as to the cortex in more basal root zones. Protein gel blot analysis of two-phase partitioned root membrane fractions and whole-mount immunolocalization in root hairs revealed the plasma membrane to be enriched in AtDUR3 protein. Expression of the AtDUR3 gene in nitrogen-deficient roots was repressed by ammonium and nitrate but induced after supply of urea. Higher accumulation of urea in roots of wild-type plants relative to atdur3-1 and atdur3-3 confirmed that urea was the substrate transported by AtDUR3. Influx of 15N-labeled urea in atdur3-1 and atdur3-3 showed a linear concentration dependency up to 200 microM external urea, whereas influx in wild-type roots followed saturation kinetics with an apparent Km of 4 microM. The results indicate that AtDUR3 is the major transporter for high-affinity urea uptake in Arabidopsis roots and suggest that the high substrate affinity of AtDUR3 reflects an adaptation to the low urea levels usually found in unfertilized soils.

The AMMONIUM TRANSPORTER (AMT) family comprises six isoforms in Arabidopsis thaliana. Here, we describe the complete functional organization of root-expressed AMTs for high-affinity ammonium uptake. High-affinity influx of (15)N-labeled ammonium in two transposon-tagged amt1;2 lines was reduced by 18 to 26% compared with wild-type plants. Enrichment of the AMT1;2 protein in the plasma membrane and localization of AMT1;2 promoter activity in the endodermis and root cortex indicated that AMT1;2 mediates the uptake of ammonium entering the root via the apoplasmic transport route. An amt1;1 amt1;2 amt1;3 amt2;1 quadruple mutant (qko) showed severe growth depression under ammonium supply and maintained only 5 to 10% of wild-type high-affinity ammonium uptake capacity. Transcriptional upregulation of AMT1;5 in nitrogen-deficient rhizodermal and root hair cells and the ability of AMT1;5 to transport ammonium in yeast suggested that AMT1;5 accounts for the remaining uptake capacity in qko. Triple and quadruple amt insertion lines revealed in vivo ammonium substrate affinities of 50, 234, 61, and 4.5 muM for AMT1;1, AMT1;2, AMT1;3, and AMT1;5, respectively, but no ammonium influx activity for AMT2;1. These data suggest that two principle means of achieving effective ammonium uptake in Arabidopsis roots are the spatial arrangement of AMT1-type ammonium transporters and the distribution of their transport capacities at different substrate affinities.

In Arabidopsis four root-expressed AMT genes encode functional ammonium transporters, which raises the question of their role in primary ammonium uptake. After pre-culturing under nitrogen-deficiency conditions, we quantified the influx of (15)N-labeled ammonium in T-DNA insertion lines and observed that the loss of either AMT1;1 or AMT1;3 led to a decrease in the high-affinity ammonium influx of approximately 30%. Under nitrogen-sufficient conditions the ammonium influx was lower in Columbia glabra compared with Wassilewskija (WS), and AMT1;1 did not contribute significantly to the ammonium influx in Col-gl. Ectopic expression of AMT1;3 under the control of a 35S promoter in either of the insertion lines amt1;3-1 or amt1;1-1 increased the ammonium influx above the level of their corresponding wild types. In transgenic lines carrying AMT-promoter-GFP constructs, the promoter activities of AMT1;1 and AMT1;3 were both upregulated under nitrogen-deficiency conditions and were localized to the rhizodermis, including root hairs. AMT gene-GFP fusions that were stably expressed under the control of their own promoters were localized to the plasma membrane. The double insertion line amt1;1-1amt1;3-1 showed a decreased sensitivity to the toxic ammonium analog methylammonium and a decrease in the ammonium influx of up to 70% relative to wild-type plants. These results suggest an additive contribution of AMT1;1 and AMT1;3 to the overall ammonium uptake capacity in Arabidopsis roots under nitrogen-deficiency conditions.

Root system architecture (RSA) is seldom considered as a selection criterion to improve yield in maize breeding, mainly because of the practical difficulties with their evaluation under field conditions. In the present study, phenotypic profiling of 187 advanced-backcross BCF maize lines (Ye478 × Wu312) was conducted at different developmental stages under field conditions at two locations (Dongbeiwang in 2007 and Shangzhuang in 2008) for five quantitative root traits. The aims were to (1) understand the genetic basis of root growth in the field; (2) investigate the contribution of root traits to grain yield (GY); and (3) detect QTLs controlling root traits at the seedling (I), silking (II) and maturation (III) stages. Axial root (AR)-related traits showed higher heritability than lateral root (LR)-related traits, which indicated stronger environmental effects on LR growth. Among the three developmental stages, root establishment at stage I showed the closest relationship with GY ( r = 0.33-0.43, P < 0.001). Thirty QTLs for RSA were detected in the BCF population and only 13.3 % of the QTLs were detected at stage III. Most important QTLs for root traits were located on chromosome 6 near the locus umc1257 (bin 6.02-6.04) at stage I, and chromosome 10 near the locus umc2003 (bin 10.04) for number of AR across all three developmental stages. The regions of chromosome 7 near the locus bnlg339 (bin 7.03) and chromosome 1 near the locus bnlg1556 (bin 1.07) harbored QTLs for both GY- and LR-related traits at stages I and II, respectively. These results help to understand the genetic basis of root development under field conditions and their contribution to grain yield. [ABSTRACT FROM AUTHOR]

In order to understand the regulation of source-sink relationship on leaf senescence in maize, we investigated the effect of pollination prevention on leaf senescence, post-silking dry matter and nitrogen accumulation and remobilization in different vegetative organs in one landrace and ten representative maize hybrids released between 1973 and 2000 in China. When pollination was prevented by covering silk with paper bags at silking stage, leaf senescence of Nongda 60, Yedan 13, Shendan 7 and Zhengdan 958 was delayed as shown by the remaining green leaf area per plant at maturity. However, pollination-prevention did not have obvious effect on the leaf senescence of Baimaya, Danyu 6, Yedan 2 and Xianyu 335. By contrast, pollination-prevention accelerated the leaf senescence of maize hybrids Zhongdan 2, Danyu13 and Yedan 4. It was found that leaf senescence of the late-released (since 1985) Chinese hybrids tended to be delayed by pollination-prevention, except XY335. From silking to physiological maturity, nitrogen content increased in the stem (plus the sheath, cob, husk, and tassel) and root of the non-pollinated plants. However, there was still a net reduction in leaf nitrogen content. We also found that pollination-prevention reduced leaf nitrogen remobilization efficiency, with genotypic difference and variation between the two years of testing. The results suggested that the response of leaf senescence to pollination-prevention is at least partially due to the change of leaf nitrogen remobilization efficiency. Leaf senescence tended to be delayed if leaf nitrogen remobilization efficiency is highly reduced by pollination-prevention. [ABSTRACT FROM AUTHOR]