Root hair phenotypes influence nitrogen acquisition in maize.

BACKGROUND AND AIMS: The utility of root hairs for nitrogen (N) acquisition is poorly understood. METHODS: We explored the utility of root hairs for N acquisition in the functional-structural model SimRoot and with maize genotypes with variable root hair length (RHL) in greenhouse and field environments. KEY RESULTS: Simulation results indicate that long, dense root hairs can improve N acquisition under varying N availability. In the greenhouse, ammonium availability had no effect on RHL and low nitrate availability increased RHL, while in the field low N reduced RHL. Longer RHL was associated with 216 % increase in biomass and 237 % increase in plant N content under low-N conditions in the greenhouse and a 250 % increase in biomass and 200 % increase in plant N content in the field compared with short-RHL phenotypes. In a low-N field environment, genotypes with long RHL had 267 % greater yield than those with short RHL. We speculate that long root hairs improve N capture by increased root surface area and expanded soil exploration beyond the N depletion zone surrounding the root surface. CONCLUSIONS: We conclude that root hairs play an important role in N acquisition. We suggest that root hairs merit consideration as a breeding target for improved N acquisition in maize and other crops.

Reduced root cortical cell file number improves drought tolerance in maize.

We tested the hypothesis that reduced root cortical cell file number (CCFN) would improve drought tolerance in maize (Zea mays) by reducing the metabolic costs of soil exploration. Maize genotypes with contrasting CCFN were grown under well-watered and water-stressed conditions in greenhouse mesocosms and in the field in the United States and Malawi. CCFN ranged from six to 19 among maize genotypes. In mesocosms, reduced CCFN was correlated with 57% reduction of root respiration per unit of root length. Under water stress in the mesocosms, genotypes with reduced CCFN had between 15% and 60% deeper rooting, 78% greater stomatal conductance, 36% greater leaf CO2 assimilation, and between 52% to 139% greater shoot biomass than genotypes with many cell files. Under water stress in the field, genotypes with reduced CCFN had between 33% and 40% deeper rooting, 28% lighter stem water oxygen isotope enrichment (δ(18)O) signature signifying deeper water capture, between 10% and 35% greater leaf relative water content, between 35% and 70% greater shoot biomass at flowering, and between 33% and 114% greater yield than genotypes with many cell files. These results support the hypothesis that reduced CCFN improves drought tolerance by reducing the metabolic costs of soil exploration, enabling deeper soil exploration, greater water acquisition, and improved growth and yield under water stress. The large genetic variation for CCFN in maize germplasm suggests that CCFN merits attention as a breeding target to improve the drought tolerance of maize and possibly other cereal crops.

Large root cortical cell size improves drought tolerance in maize.

The objective of this study was to test the hypothesis that large cortical cell size (CCS) would improve drought tolerance by reducing root metabolic costs. Maize (Zea mays) lines contrasting in root CCS measured as cross-sectional area were grown under well-watered and water-stressed conditions in greenhouse mesocosms and in the field in the United States and Malawi. CCS varied among genotypes, ranging from 101 to 533 µm(2). In mesocosms, large CCS reduced respiration per unit of root length by 59%. Under water stress in mesocosms, lines with large CCS had between 21% and 27% deeper rooting (depth above which 95% of total root length is located in the soil profile), 50% greater stomatal conductance, 59% greater leaf CO2 assimilation, and between 34% and 44% greater shoot biomass than lines with small CCS. Under water stress in the field, lines with large CCS had between 32% and 41% deeper rooting (depth above which 95% of total root length is located in the soil profile), 32% lighter stem water isotopic ratio of (18)O to (16)O signature, signifying deeper water capture, between 22% and 30% greater leaf relative water content, between 51% and 100% greater shoot biomass at flowering, and between 99% and 145% greater yield than lines with small cells. Our results are consistent with the hypothesis that large CCS improves drought tolerance by reducing the metabolic cost of soil exploration, enabling deeper soil exploration, greater water acquisition, and improved growth and yield under water stress. These results, coupled with the substantial genetic variation for CCS in diverse maize germplasm, suggest that CCS merits attention as a potential breeding target to improve the drought tolerance of maize and possibly other cereal crops.

Root cortical aerenchyma enhances nitrogen acquisition from low-nitrogen soils in maize.

Suboptimal nitrogen (N) availability is a primary constraint for crop production in developing nations, while in rich nations, intensive N fertilization carries substantial environmental and economic costs. Therefore, understanding root phenes that enhance N acquisition is of considerable importance. Structural-functional modeling predicts that root cortical aerenchyma (RCA) could improve N acquisition in maize (Zea mays). We evaluated the utility of RCA for N acquisition by physiological comparison of maize recombinant inbred lines contrasting in RCA grown under suboptimal and adequate N availability in greenhouse mesocosms and in the field in the United States and South Africa. N stress increased RCA formation by 200% in mesocosms and by 90% to 100% in the field. RCA formation substantially reduced root respiration and root N content. Under low-N conditions, RCA formation increased rooting depth by 15% to 31%, increased leaf N content by 28% to 81%, increased leaf chlorophyll content by 22%, increased leaf CO2 assimilation by 22%, increased vegetative biomass by 31% to 66%, and increased grain yield by 58%. Our results are consistent with the hypothesis that RCA improves plant growth under N-limiting conditions by decreasing root metabolic costs, thereby enhancing soil exploration and N acquisition in deep soil strata. Although potential fitness tradeoffs of RCA formation are poorly understood, increased RCA formation appears be a promising breeding target for enhancing crop N acquisition.

Root anatomical phenes predict root penetration ability and biomechanical properties in maize (Zea Mays).

The ability of roots to penetrate hard soil is important for crop productivity but specific root phenes contributing to this ability are poorly understood. Root penetrability and biomechanical properties are likely to vary in the root system dependent on anatomical structure. No information is available to date on the influence of root anatomical phenes on root penetrability and biomechanics. Root penetration ability was evaluated using a wax layer system. Root tensile and bending strength were evaluated in plant roots grown in the greenhouse and in the field. Root anatomical phenes were found to be better predictors of root penetrability than root diameter per se and associated with smaller distal cortical region cell size. Smaller outer cortical region cells play an important role in stabilizing the root against ovalization and reducing the risk of local buckling and collapse during penetration, thereby increasing root penetration of hard layers. The use of stele diameter was found to be a better predictor of root tensile strength than root diameter. Cortical thickness, cortical cell count, cortical cell wall area and distal cortical cell size were stronger predictors of root bend strength than root diameter. Our results indicate that root anatomical phenes are important predictors for root penetrability of high-strength layers and root biomechanical properties.

Root anatomical phenes associated with water acquisition from drying soil: targets for crop improvement.

Several root anatomical phenes affect water acquisition from drying soil, and may therefore have utility in breeding more drought-tolerant crops. Anatomical phenes that reduce the metabolic cost of the root cortex ('cortical burden') improve soil exploration and therefore water acquisition from drying soil. The best evidence for this is for root cortical aerenchyma; cortical cell file number and cortical senescence may also be useful in this context. Variation in the number and diameter of xylem vessels strongly affects axial water conductance. Reduced axial conductance may be useful in conserving soil water so that a crop may complete its life cycle under terminal drought. Variation in the suberization and lignification of the endodermis and exodermis affects radial water conductance, and may therefore be important in reducing water loss from mature roots into dry soil. Rhizosheaths may protect the water status of young root tissue. Root hairs and larger diameter root tips improve root penetration of hard, drying soil. Many of these phenes show substantial genotypic variation. The utility of these phenes for water acquisition has only rarely been validated, and may have strong interactions with the spatiotemporal dynamics of soil water availability, and with root architecture and other aspects of the root phenotype. This complexity calls for structural-functional plant modelling and 3D imaging methods. Root anatomical phenes represent a promising yet underexplored and untapped source of crop breeding targets.

Root cortical burden influences drought tolerance in maize.

BACKGROUND AND AIMS: Root cortical aerenchyma (RCA) increases water and nutrient acquisition by reducing the metabolic costs of soil exploration. In this study the hypothesis was tested that living cortical area (LCA; transversal root cortical area minus aerenchyma area and intercellular air space) is a better predictor of root respiration, soil exploration and, therefore, drought tolerance than RCA formation or root diameter. METHODS: RCA, LCA, root respiration, root length and biomass loss in response to drought were evaluated in maize (Zea mays) recombinant inbred lines grown with adequate and suboptimal irrigation in soil mesocosms. KEY RESULTS: Root respiration was highly correlated with LCA. LCA was a better predictor of root respiration than either RCA or root diameter. RCA reduced respiration of large-diameter roots. Since RCA and LCA varied in different parts of the root system, the effects of RCA and LCA on root length were complex. Greater crown-root LCA was associated with reduced crown-root length relative to total root length. Reduced LCA was associated with improved drought tolerance. CONCLUSIONS: The results are consistent with the hypothesis that LCA is a driver of root metabolic costs and may therefore have adaptive significance for water acquisition in drying soil.

Agronomic biofortification increases grain zinc concentration of maize grown under contrasting soil types in Malawi.

Zinc (Zn) deficiency remains a public health problem in Malawi, especially among poor and marginalized rural populations, linked with low dietary intake of Zn due to consumption of staple foods that are low in Zn content. The concentration of Zn in staple cereal grain can be increased through application of Zn-enriched fertilizers, a process called agronomic biofortification or agro-fortification. Field experiments were conducted at three Agricultural Research Station sites to assess the potential of agronomic biofortification to improve Zn concentration in maize grain in Malawi as described in registered report published previously. The hypotheses of the study were (i) that application of Zn-enriched fertilizers would increase in the concentration of Zn in maize grain to benefit dietary requirements of Zn and (ii) that Zn concentration in maize grain and the effectiveness of agronomic biofortification would be different between soil types. At each site two different subsites were used, each corresponding to one of two agriculturally important soil types of Malawi, Lixisols and Vertisols. Within each subsite, three Zn fertilizer rates (1, 30, and 90 kg ha-1) were applied to experimental plots, using standard soil application methods, in a randomized complete block design. The experiment had 10 replicates at each of the three sites as informed by a power analysis from a pilot study, published in the registered report for this experiment, designed to detect a 10% increase in grain Zn concentration at 90 kg ha-1, relative to the concentration at 1 kg ha-1. At harvest, maize grain yield and Zn concentration in grain were measured, and Zn uptake by maize grain and Zn harvest index were calculated. At 30 kg ha-1, Zn fertilizer increased maize grain yields by 11% compared with nationally recommended application rate of 1 kg ha-1. Grain Zn concentration increased by 15% and uptake by 23% at the application rate of 30 kg ha-1 relative to the national recommendation rate. The effects of Zn fertilizer application rate on the response variables were not dependent on soil type. The current study demonstrates the importance of increasing the national recommendation rate of Zn fertilizer to improve maize yield and increase the Zn nutritional value of the staple crop.

Increasing zinc concentration in maize grown under contrasting soil types in Malawi through agronomic biofortification: Trial protocol for a field experiment to detect small effect sizes.

The prevalence of micronutrient deficiencies including zinc (Zn) is widespread in Malawi, especially among poor and marginalized rural populations. This is due to low concentrations of Zn in most staple cereal crops and limited consumption of animal source foods. The Zn concentration of cereal grain can be increased through application of Zn-enriched fertilizers; a process termed agronomic biofortification or agro-fortification. This trial protocol describes a field experiment which aims to assess the potential of agronomic biofortification to improve the grain Zn concentration of maize, the predominant staple crop of Malawi. The hypotheses of the study are that application of Zn-enriched fertilizers will create a relatively small increase in the concentration of Zn in maize grains that will be sufficient to benefit dietary supplies of Zn, and that the effectiveness of agronomic biofortification will differ between soil types. The study will be conducted at three sites, Chitedze, Chitala, and Ngabu Agricultural Research Stations, in Lilongwe, Salima, and Chikwawa Districts respectively. These three sites represent locations in the Central and Southern Regions of Malawi. At each site, two different sub-sites will be used, each corresponding to one of two agriculturally important soil types of Malawi, Lixisols, and Vertisols. Within each sub-site, three Zn fertilizer rates (1, 30, and 90 kg/ha) will be applied to experimental plots using standard soil application methods, in a randomized complete block design. The number of replicates at plot level has been informed by a power analysis from pilot study data, assuming that a minimum 10% increase in Zn concentration of grain at 90 kg/ha relative to the concentration at 1 kg/ha is of interest. Grain mass (yield), stover mass, and both stover and grain Zn concentrations will be measured at harvest. A second year of cropping will be used to establish whether there are any residual benefits to grain Zn concentration. The potential for Zn agronomic biofortification will be communicated to relevant academic and government stakeholders through a peer review journal article and a briefing paper.