Understanding Root Systems to Improve Seedling Quality

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.

Root systems play a pivotal role in water and nutrient uptake from soil. Lateral root (LR) growth is promoted to compensate for inhibited main root growth. Compensatory LR growth contributes to maintaining total root length (TRL) and hence water and nutrient uptake in compacted soils. However, it remains unclear how shoot and root phenotypic traits change during the compensatory growth and whether there are genotypic variations in compensatory root growth. This study analyzed shoot and root morphological traits of 20 rice genotypes, which includes mutants with altered root morphology, during the vegetative stage using a semihydroponic phenotyping system. The phenotyping experiment detected large variation in root and shoot traits among the 20 genotypes. Morphological changes induced by root cutting were analyzed in six selected genotypes with contrasting root system architecture. Root cutting significantly affected root distribution along vertical sections and among diameter classes. After root cutting, more roots distributed at shallower depth and thicker LRs developed. Furthermore, genotypes with deeper root growth without root cutting allocated more compensatory roots to deeper sections even after root cutting than the genotypes with shallower rooting. Due to the compensatory LR growth, root cutting did not significantly affect TRL, root dry weight (RDW), or shoot dry weight (SDW). To analyze the interaction between crown root (CR) number and compensatory root growth, we removed half of the newly emerged CRs in two genotypes. TRL of YRL38 increased at depth with CR number manipulation (CRM) regardless of root tip excision, which was attributed to an increase in specific root length (SRL), despite no change in RDW. Taken together, the tested rice genotypes exhibited compensatory root growth by changing root distribution at depth and in diameter classes. Reducing CR number promoted root development and compensatory growth by improving the efficiency of root development [root length (RL) per resource investment].

ABSTRACTThe rice root system consists of three types of roots; main root, L-type and S-type lateral root (LR). These component roots are morphologically and histologically different, which is termed as heterorhizy. Root system hydraulic architecture is related to the unique features of component roots. We hypothesized that each component root contributes in different degrees to water uptake of the whole root system. Rice varieties IRAT 109 and Taichung 65 were grown in pots filled with soil under continuous waterlogged (CWL) and drought (CD) conditions until two weeks after heading. Morphology and histological structures of roots, which may regulate radial water movement, were compared among the three component roots. Moreover, hydraulic conductivity (Lpr) of the root system, which represents the water uptake ability, were measured with a pressure chamber. Based on a model that Lpr of the whole root system is a product of Lpr of each of the component roots and their surface areas, we found that the differences in Lpr between the two varieties and the plants grown under different soil water conditions for any of the component roots did not support the corresponding differences in the measured Lpr of the whole root system. In contrast, a significant and positive correlation was found between Lpr of the whole root system and the percentage of surface area of S-type LR but not for the other component roots. These results indicate S-type LR might have a higher contribution to Lpr of the whole root system than the other component roots.

Barley (Hordeum vulgare L) may alter its root morphology to improve P acquisition efficiency under low-P (LP) stress. This research studied the variations in adventitious and lateral root morphological traits of barley and mapped their quantitative trait loci (QTLs) under LP and high P (HP). The recombinant inbred lines were derived from the F1 population of a cross between CN4027 and Baudin. Two experiments aimed to identify QTLs related to adventitious and lateral root morphological traits under LP and HP. The length, surface area and volume of adventitious and lateral roots were measured. Under HP, Baudin had larger root morphology, especially lateral root morphology, than CN4027. LP stress induced lateral root growth but inhibited adventitious root growth. Nineteen QTLs for root morphological traits were detected. These QTLs clustered within four regions (Cl-2H, Cl-3H, Cl-4H and Cl-7H) on chromosomes 2H, 3H, 4H and 7H, with corresponding contributions of 12.0-42.9%. Some QTLs are linked with the QTLs for P efficiency detected previously, demonstrating the role of root morphological traits in P efficiency. The Cl-2H region was identified in the interval bPb3927665-bPb3265744 on chromosome 2H and had major effects on lateral root growth, especially under LP. Lateral root length and surface area increased when alleles from Baudin were present at the QTLs in Cl-2H. This study demonstrated the patterns of growth among root types and the role of lateral roots in barley's adaption to LP stress. The QTL clusters, especially Cl-2H, may offer clues for fine mapping and map-based cloning.

The root system commonly lies underground, where it provides anchorage for the aerial organs, as well as nutrients and water. Both endogenous and environmental cues contribute to the establishment of the root system. Among the endogenous cues, microRNAs (miRNAs), transcription factors, and phytohormones modulate root architecture. miRNAs belong to a subset of endogenous hairpin-derived small RNAs that post-transcriptionally control target gene expression, mostly transcription factors, comprising the miRNA regulatory hubs. Phytohormones are signaling molecules involved in most developmental processes. Some miRNAs and targets participate in more than one hormonal pathway, thereby providing new bridges in plant hormonal crosstalk. Unraveling the intricate network of molecular mechanisms underlying the establishment of root systems is a central aspect in the development of novel strategies for plant breeding to increase yield and optimize agricultural land use. In this review, we summarize recent findings describing the molecular mechanisms associated with the interplay between miRNA regulatory hubs and phytohormones to ensure the establishment of a proper root system. We focus on post-embryonic growth and development of primary, lateral, and adventitious roots. In addition, we discuss novel insights for future research on the interaction between miRNAs and phytohormones in root architecture.

Understanding the development of roots exposed to contaminants and the potential of plant-associated bacteria for optimization of growth.

Lupinus angustifolius L. (narrow-leafed lupin) is an important grain legume crop for the stockfeed industry in Australia. This species does not form cluster roots regardless of phosphorus (P) nutrition. We hypothesise that this species may have adaptive strategies for achieving critical P uptake in low-P environments by altering shoot growth and root architecture and secreting carboxylates from roots. Three wild genotypes of L. angustifolius with contrasting root architecture were selected to investigate the influence of P starvation on root growth and rhizosphere carboxylate exudation and their relationship with P acquisition. Plants were grown in sterilised loamy soil supplied with zero, low (50 µm) or optimal (400 µm) P for 6 weeks. All genotypes showed a significant response in shoot and root development to varying P supply. At P deficit (zero and low P), root systems were smaller and had fewer branches than did roots at optimal P. The amount of total carboxylates in the rhizosphere extracts ranged from 3.4 to 17.3 µmol g–1 dry root. The total carboxylates comprised primarily citrate (61–78% in various P treatments), followed by malate and acetate. Genotype #085 (large root system with deep lateral roots) exuded the greatest amount of total carboxylates to the rhizosphere for each P treatment, followed by #016 (medium root system with good branched lateral roots) and #044 (small root system with short and sparse lateral roots). All genotypes in the low-P treatment significantly enhanced exudation of carboxylates, whereas no significant increase in carboxylate exudation was observed in the zero-P treatment. Small-rooted genotypes had higher P concentration than the medium- and large-rooted genotypes, although larger plants accumulated higher total P content. Large-rooted genotypes increased shoot P utilisation efficiency in response to P starvation. This study showed that narrow-leafed lupin genotypes differing in root architecture differed in carboxylate exudation and P uptake. Our finding suggested that for L. angustifolius there is a minimum plant P concentration below which carboxylate exudation is not enhanced despite severe P deficiency. The outcomes of this study enhance our understanding of P acquisition strategies in L. angustifolius genotypes, which can be used for the selection of P-efficient genotypes for cropping systems.

Determining the variation in roots traits within a grass root system is important for understanding the role of fine roots in carbon and nutrient cycling in grassland ecosystems, where the majority of biomass and litter accumulation occur belowground. However, few studies have been conducted in this regard. In this study, the structural and chemical traits of shoot-derived and root-derived roots were examined in three perennial grasses—Cleistogenes squarrosa, Achnatherum sibiricum and Stipa grandis—aiming to explore structural differences, responses to nitrogen and water addition in different types of roots and their correlations with aboveground plant nitrogen. Our results showed significant differences between these two root types, with root-derived roots having higher N concentration, tissue density, and specific root length, but lower C: N and diameter than shoot-derived roots. Trait relationships between root N concentration and tissue density for the two root types differed from that reported among species. These traits in different types of roots were insensitive to resource addition. Furthermore, N concentration in shoot-derived roots was more strongly linked to aboveground plant N concentration than root-derived roots. The results of this study demonstrate structural differences within the root system that may reflect functional heterogeneity in grass roots.

Lateral root development is an important morphogenetic process in plants, which allows the modulation root architecture and substantially determines the plant's efficiency for water and nutrient uptake. Postembryonic root development is under the control of both endogenous developmental programs and environmental stimuli. Nutrient availability plays a major role among environmental signals that modulate root development. Phosphate (Pi) limitation is a constraint for plant growth in many natural and agricultural ecosystems. Plants posses Pi-sensing mechanisms that enable them to respond and adapt to conditions of limited Pi supply, including increased formation and growth of lateral roots. Root developmental modifications are mainly mediated by the plant hormone auxin. Recently we showed that the alteration of root system architecture under Pi-starvation may be mediated by modifications in auxin sensitivity in root cells via a mechanism involving the TIR1 auxin receptor. In this addendum, we provide additional novel evidence indicating that the low Pi pathway involves changes in cell cycle gene expression. It was found that Pi deprivation increases the expression of CDKA, E2Fa, Dp-E2F and CyCD3. In particular, E2Fa, Dp-E2F and CyCD3 genes were specifically up-regulated by auxin in Pi-deprived Arabidopsis seedlings that were treated with the auxin transport inhibitor NPA, indicating that cell cycle modulation by low Pi signaling is independent of auxin transport and dependent on auxin sensitivity in the root.

The promoted lateral root 1 (plr1) mutation is involved in reduced basal shoot starch accumulation and increased root sugars for enhanced lateral root growth in rice

Maize genotypes may adapt to dry environments by avoiding desiccation by means of a deeper root system or by maintaining growth and water extraction at low water potentials. The aim of this study was to determine the quantitative genetic control of root growth and root morphology in a population of 236 recombinant inbred lines (RILs) from the cross between CML444 (high-yielding) x SC-Malawi (low-yielding), which segregates for the response to drought stress at flowering. The RILs and the parental lines were grown on blotting paper in growth pouches until the two-leaf stage under non-stressed conditions; the parents were additionally exposed to desiccation stress induced by polyethylene glycol with a molecular weight of 8000 Dalton (PEG-8000). The lengths of axile and lateral roots were measured non-destructively at 2, 5, 7 and 9 days after germination, by scanning with an A4 scanner followed by digital image analysis. CML444 had a lower rate constant of lateral root elongation (k(Lat)) than SC-Malawi, but the two genotypes did not differ in their response to desiccation. QTLs affecting root vigor, as depicted by increments in k(Lat), the elongation rate of axile roots (ER(Ax)) and the number of axile roots (No(Ax)) were identified in bins 2.04 and 2.05. QTLs for No(Ax) and ER(Ax) collocated with QTLs for yield parameters in bins 1.03-1.04 and 7.03-04. The correspondence of QTLs for axile root traits in bins 1.02-1.03 and 1.08 and QTLs for lateral roots traits in bins 2.04-2.07 in several mapping populations suggests the presence of genes controlling root growth in a wide range of genetic backgrounds.

The core regulators which are required for primary root growth and development also function in lateral root development or lateral root stem cell niche maintenance. The primary root systems and the lateral root systems are the two important root systems which are vital to the survival of plants. Though the molecular mechanism of the growth and development of both the primary root systems and the lateral root systems have been extensively studied individually in Arabidopsis, there are not so much evidence to show that if both root systems share common regulatory mechanisms. AP2 family transcription factors such as PLT1 (PLETHORA1) and PLT2, GRAS family transcription factors such as SCR (SCARECROW) and SHR (SHORT ROOT) and WUSCHEL-RELATED HOMEOBOX transcription factor WOX5 have been extensively studied and found to be essential for primary root growth and development. In this study, through the expression pattern analysis and mutant examinations, we found that these core regulators also function in lateral root development or lateral root stem cell niche maintenance.

BackgroundArabidopsis root growth is stimulated by Piriformospora indica, phosphate limitation and inactivation of the WRKY6 transcription factor. Combinations of these factors induce unexpected alterations in root and shoot growth, root architecture and root gene expression profiles.ResultsThe results demonstrate that P. indica promotes phosphate uptake and root development under Pi limitation in wrky6 mutant. This is associated with the stimulation of PHOSPHATE1 expression and ethylene production. Expression profiles from the roots of wrky6 seedlings identified genes involved in hormone metabolism, transport, meristem, cell and plastid proliferation, and growth regulation. 25 miRNAs were also up-regulated in these roots. We generated and discuss here a list of common genes which are regulated in growing roots and which are common to all three growth stimuli investigated in this study.ConclusionSince root development of wrky6 plants exposed to P. indica under phosphate limitation is strongly promoted, we propose that common genes which respond to all three growth stimuli are central for the control of root growth and architecture. They can be tested for optimizing root growth in model and agricultural plants.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-015-0673-4) contains supplementary material, which is available to authorized users.

Root architecture is key in determining how effective plants are at intercepting and absorbing nutrients and water. Previously, the wheat (Triticum aestivum) cultivars Spica and Maringa were shown to have contrasting root morphologies. These cultivars were crossed to generate an F6:1 population of recombinant inbred lines (RILs) which was genotyped using a 90 K single nucleotide polymorphisms (SNP) chip. A total of 227 recombinant inbred lines (RILs) were grown in soil for 21 days in replicated trials under controlled conditions. At harvest, the plants were scored for seven root traits and two shoot traits. An average of 7.5 quantitative trait loci (QTL) were associated with each trait and, for each of these, physical locations of the flanking markers were identified using the Chinese Spring reference genome. We also compiled a list of genes from wheat and other monocotyledons that have previously been associated with root growth and morphology to determine their physical locations on the Chinese Spring reference genome. This allowed us to determine whether the QTL discovered in our study encompassed genes previously associated with root morphology in wheat or other monocotyledons. Furthermore, it allowed us to establish if the QTL were co-located with the QTL identified from previously published studies. The parental lines together with the genetic markers generated here will enable specific root traits to be introgressed into elite wheat lines. Moreover, the comprehensive list of genes associated with root development, and their physical locations, will be a useful resource for researchers investigating the genetics of root morphology in cereals.

Circadian clocks synchronized with the environment allow plants to anticipate recurring daily changes and give a fitness advantage. Here, we mapped the dynamic growth phenotype of leaves and roots in two lines of Arabidopsis thaliana with a disrupted circadian clock: the CCA1 over-expressing line (CCA1ox) and the prr9 prr7 prr5 (prr975) mutant. We demonstrate leaf growth defects due to a disrupted circadian clock over a 24 h time scale. Both lines showed enhanced leaf growth compared with the wild-type during the diurnal period, suggesting increased partitioning of photosynthates for leaf growth. Nocturnal leaf growth was reduced and growth inhibition occurred by dawn, which may be explained by ineffective starch degradation in the leaves of the mutants. However, this growth inhibition was not caused by starch exhaustion. Overall, these results are consistent with the notion that the defective clock affects carbon and energy allocation, thereby reducing growth capacity during the night. Furthermore, rosette morphology and size as well as root architecture were strikingly altered by the defective clock control. Separate analysis of the primary root and lateral roots revealed strong suppression of lateral root formation in both CCA1ox and prr975, accompanied by unusual changes in lateral root growth direction under light-dark cycles and increased lateral extension of the root system. We conclude that growth of the whole plant is severely affected by improper clock regulation in A.thaliana, resulting not only in altered timing and capacity for growth but also aberrant development of shoot and root architecture.