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Abstract Background Root system architecture (RSA) is an essential characteristic for efficient water and nutrient absorption in terrestrial plants; its plasticity enables plants to respond to different soil environments. Better understanding of root plasticity is important in developing stress-tolerant crops. Non-invasive techniques that can measure roots in soils nondestructively, such as X-ray computed tomography (CT), are useful to evaluate RSA plasticity. However, although RSA plasticity can be measured by tracking individual root growth, only a few methods are available for tracking individual roots from time-series three-dimensional (3D) images. Results We developed a semi-automatic workflow that tracks individual root growth by vectorizing RSA from time-series 3D images via two major steps. The first step involves 3D alignment of the time-series RSA images by iterative closest point registration with point clouds generated by high-intensity particles in potted soils. This alignment ensures that the time-series RSA images overlap. The second step consists of backward prediction of vectorization, which is based on the phenomenon that the root length of the RSA vector at the earlier time point is shorter than that at the last time point. In other words, when CT scanning is performed at time point A and again at time point B for the same pot, the CT data and RSA vectors at time points A and B will almost overlap, but not where the roots have grown. We assumed that given a manually created RSA vector at the last time point of the time series, all RSA vectors except those at the last time point could be automatically predicted by referring to the corresponding RSA images. Using 21 time-series CT volumes of a potted plant of upland rice (Oryza sativa), this workflow revealed that the root elongation speed increased with age. Compared with a workflow that does not use backward prediction, the workflow with backward prediction reduced the manual labor time by 95%. Conclusions We developed a workflow to efficiently generate time-series RSA vectors from time-series X-ray CT volumes. We named this workflow 'RSAtrace4D' and are confident that it can be applied to the time-series analysis of RSA development and plasticity.

Changes in root development and growth in upland rice receiving subsurface irrigation were evaluated using root box experiments. Irrigated water was independently applied every day to depths of 0, 5, 10 and 20 cm from the ground surface, and root morphology and aboveground growth were determined after 2 weeks of treatment. Plant length, stem number and leaf age significantly increased along with the irrigation depth, and the aboveground dry weights of rice grown at 5-, 10- and 20-cm depths were significantly greater than those grown at 0 cm. Root surface area significantly increased along with the irrigation depth, which was attributed to the increasing lengths and branch numbers of roots. Subsurface irrigation significantly increased root lengths in deep layers (below 15 cm) and increased the deep layers’ distribution ratios by 1.2 to 5.7 times as the surface layer ratio decreased. This was accompanied with a decrease in the root diameter. This may be associated with the increase in the numbers of branches and the decrease in the numbers of thick nodal roots. These results indicate that subsurface irrigation increases root length and branching in the deep soil layers, inducing deep rooting, even with same amount of irrigation.

Centipedegrass [Eremochloa ophiuroides (Munro) Hack.] is a perennial warm-season grass that originated in China, and its speed of nodal rooting is important for lawn establishment. In our study, centipedegrass nodal rooting ability was limited by node aging. Transcriptome sequencing of nodal roots after 0, 2, 4, and 8 days of water culture was performed to investigate the molecular mechanisms of root development. GO enrichment and KEGG pathway analyses of DEGs indicated that plant hormone signal transduction and transcription factors might play important roles in centipedegrass nodal root growth. Among them, E3 ubiquitin-protein ligases participated in multiple hormone signal transduction pathways and interacted with transcription factors. Furthermore, an E3 ubiquitin protein ligase EoSINAT5 overexpressed in rice resulted in longer roots and more numerous root tips, while knockout of LOC_Os07g46560 (the homologous gene of EoSINAT5 in rice) resulted in shorter roots and fewer root tips. These results indicated that EoSINAT5 and its homologous gene are able to promote nodal root development. This research presents the transcriptomic analyses of centipedegrass nodal roots, and may contribute to elucidating the mechanism governing the development of nodal roots and facilitates the use of molecular breeding in improving rooting ability.

The root of wheat consists of seminal and nodal roots. Comparatively speaking, fewer studies have been carried out on the nodal root system because of its disappearance at the early seedling stage under indoor environments. In this study, 196 accessions from the Huanghuai Wheat Region (HWR) were used to identify the characteristics of seminal and nodal root traits under different growth environments, including indoor hydroponic culture (IHC), outdoor hydroponic culture (OHC), and outdoor pot culture (OPC), for three growing seasons. The results indicated that the variation range of root traits in pot environment was larger than that in hydroponic environment, and canonical coefficients were the greatest between OHC and OPC (0.86) than those in other two groups, namely, IHC vs. OPC (0.48) and IHC vs. OHC (0.46). Most root traits were negatively correlated with spikes per area (SPA), grains per spike (GPS), and grain yield (GY), while all the seminal root traits were positively correlated with thousand-kernel weight (TKW). Genome-wide association study (GWAS) was carried out on root traits by using a wheat 660K SNP array. A total of 35 quantitative trait loci (QTLs)/chromosomal segments associated with root traits were identified under OPC and OHC. In detail, 11 and 24 QTLs were significantly associated with seminal root and nodal root traits, respectively. Moreover, 13 QTLs for number of nodal roots per plant (NRP) containing 14 stable SNPs, were distributed on chromosomes 1B, 2B, 3A, 4B, 5D, 6D, 7A, 7B, and Un. Based on LD and bioinformatics analysis, these QTLs may contain 17 genes closely related to NRP. Among them, TraesCS2B02G552500 and TraesCS7A02G428300 were highly expressed in root tissues. Moreover, the frequencies of favorable alleles of these 14 SNPs were confirmed to be less than 70% in the natural population, suggesting that the utilization of these superior genes in wheat root is still improving.

Deep rooting of wheat has been suggested that it influences the tolerance to various environmental stresses. In this study, the nodes from which the deepest penetrated roots had emerged were examined in winter wheat. The wheat was grown in long tubes with or without mechanical stress and in large root boxes. The length and growth angle of each axile root were examined to analyze the difference in the vertical distribution of the roots between the two wheat cultivars, one with a deep and one with a shallow root system. In Shiroganekomugi, a Japanese winter wheat cultivar with a shallow root system, the rooting depths of the seminal and nodal roots decreased as the rooting nodes advanced acropetally. Six out of nine deepest roots were seminal root in the non-mechanical stress conditions. In Mutsubenkei, a Japanese winter wheat cultivar with a deep root system, grown in root boxes, not only the seminal roots but also the coleoptilar and the first nodal roots penetrated to a depth of more than 1.3 m in the root box, and became the deepest roots. In both cultivars, the seminal roots became the deepest roots under the mechanical stress conditions. There were no clear tendencies in the root growth angles among the rooting nodes in the wheat root system. This indicates that the length of the axile roots can explain the differences in the rooting depths among axile roots in a wheat root system. On the other hand, the axile roots of Mutsubenkei elongated significantly more vertically than those of Shiroganekomugi. This suggests that not only seminal but also nodal roots exhibit strong positive gravitropism and penetrate deeply in a cultivar with a deep root system. In wheat cultivars, it is likely that the extent of its Root Depth Index results partly from the gravitropic responses of both seminal and nodal roots.

Nodal root anatomy was compared among twelve upland and lowland rice (Oryza sative L.) varieties with tropical origin which were grown in hydroponic culture and under field conditions. The traditional upland japonica varieties showed the largest diameter of root, stele, and xylem vessel followed by modern upland varieties. There was a clear varietal difference in the ratio of stele to root diameter, which was associated with the genetic group rather than with the ecosystems. The japonica varieties had a significantly larger stele diameter relative to the root diameter than indica and aus varieties. The indica and aus varieties displayed more xylem vessels per unit area of stele than the japonica varieties, but the diameter of xylem vessel was smaller. Equivalent xylem vessel diameter (De) was more dependent on the number of xylem in the indica varieties than in the japonica varieties. Distinctly different types of sclerenchyma anatomy were identified among the varieties. The development of sclerenchyma was classified into four different types based on thickening of cell wall in the outer cortical parenchyma and the number of sclerenchymatous cell layers. Like the xylem anatomy, the varietal differences in sclerenchyma development were more associated with genetic group rather than the ecotype. The japonica varieties had higher frequency of the types which have a doubled cell layer in sclerenchyma with thick cell wall than indica and aus. The difference among the genetic groups was nearly consistent across growing conditions, aerobic and submerged soils. These results indicated that sclerenchyma development is controlled by a genetic factor.

Rice plants in the rainfed areas are mostly grown under fluctuating soil moisture. We examined responses in dry matter production, root development and water use to changing soil moisture in diverse rice cultivars. Rice plants were grown in polyvinyl chloride tubes under glasshouse conditions. Progressive drought right after planting greatly inhibited the shoot dry matter production, tiller development, nodal root development and water uptake in all cultivars tested. When the plants experienced soil submergence before being exposed to drought, all the cultivars exhibited higher dry matter production than their well-watered counterparts. Cultivar differences were clearly noted in the growth responses to rewatering after these plants were droughted. With well–watered control as basis, IRAT 109 and KDML 105 plants increased efficiency in converting available dry matter to increase their total root length by means of enhanced lateral root development. In the latter, however, the dry weight of roots also increased and so did root water uptake. In Dular, droughted plants did not show a clear response in terms of root development and water uptake to rewatering while its shoot growth was much more severely inhibited than the other cultivars. These findings suggest that phenotypic plasticity in the root system structure exhibited by promoted lateral root development and new nodal root production play a key role in the growth of rice under changing moisture level in the soil.