Root Elongation Rate Research Articles

Simultaneous Overexpression of Citrate Synthase and Phosphoenolpyruvate Carboxylase in Leaves Augments Citrate Exclusion and Al Resistance in Transgenic Tobacco

Phosphoenolpyruvate carboxylase (PEPC) and citrate synthase (CS) are two key enzymes in organic acid synthesis metabolism. In the present study, a cytoplasmic form of CS from tobacco and a mutant (with reduced sensitivity to organic acid inhibition) PEPC from Synechococcus vulcanus were overexpressed simultaneously using a light-inducible promoter in tobacco leaves. The analysis for enzyme activity showed that CS and PEPC enzyme activities were increased by 235% to 257% and 218% to 236% in the selected cs and pepc (double-gene) overexpression lines, respectively, compared with those in the wild-type plants (WT). The measurement for the relative root elongation rate of the tobacco plants exposed to 30 μM aluminum (Al) indicated that Al tolerance in the double-gene overexpression lines was stronger than that of the transgenic cs or pepc lines and WT plants. The 13C-NMR analysis with NaH13CO3 showed that overexpression of CS and PEPC in the transgenic tobacco successfully constructed a new citrate synthesis pathway. Under the conditions with Al stress, the amount of citrate secreted from the double-transgenic tobacco roots was the largest among the tested plants. When grown on sandy soil supplied with a nutritional solution containing 500 μM Al, the growth of the double-transgenic tobacco was better than that of the transgenic cs or pepc tobacco and WT, and their root biomass was the highest among the tested plants. These results demonstrated that construction of a new citrate synthesis pathway by simultaneous overexpression of CS and PEPC in the cytoplasm of transgenic plant leaves could enhance Al resistance in plants.

Novel QTLs for growth angle of seminal roots in wheat (Triticum aestivum L.)

Because plants cannot change their environmental circumstances by changing their location, they must instead adapt to a wide variety of environmental conditions, especially soil conditions. One of the most effective ways for a plant to adapt to a given soil condition is by modifying its root system architecture. We aim to identify the genetic factors controlling root growth angle, a trait that affects root system architecture. The present study consisted of a genetic analysis of the seminal root growth angle in wheat; the parental varieties of the doubled haploid lines (DHLs) used in this study exhibited significantly different root growth directions. Using the ‘basket’ method, the ratio of deep roots (DRR; the proportion of total roots with GA > 45 degrees) was observed for evaluating deep rooting. We were able to identify novel quantitative trait loci (QTLs) controlling the gravitropic and hydrotropic responses of wheat roots. Moreover, we detected one QTL for seminal root number per seedling (RN) on chromosome 5A and two QTLs for seminal root elongation rate (ER) on chromosomes 5D and 7D. Gravitropic and hydrotropic responses of wheat roots, which play a significant role in establishing root system architecture, are controlled by independent genetic factors.

Joint toxic effects of heavy metals and atrazine on invasive plant species Solidago Canadensis L.

In this study, the joint effects of Cd(II), Pb(II) and atrazine (ATR) on Solidago Canadensis L. were investigated. The results showed that soil containing Cd, Pb and ATR could inhibit root elongation of Solidago Canadensis L., and that there was a positive linear relationship between the inhibitory rate of root elongation and the concentrations of Pb(II). The mixture of Cd(II) and Pb(II)-ATR in soil showed a significant adverse effect on root and shoot biomass of Solidago Canadensis L. Acetylsalicylic acid (ASA) contents increased slightly at the lower concentrations of Cd(II)-ATR, then decreased when Cd(II)-ATR concentrations were higher than those of 1.0 TUmix(Cd+ATR). The toxic effect of Pb(II)-ATR on the ASA contents of Solidago Canadensis L. was greater than that of Cd(II)-ATR. Soluble sugar contents firstly decreased and then increased with increasing concentrations of Cd(II)-ATR and displayed fluctuation with increasing concentrations of Pb(II)-ATR. Total protein contents increased with increasing concentrations of Cd(II) and Pb(II)-ATR, relative to the control sample. The NP-SH contents showed a significant increase up to 12.2 mg/gfw of Cd(II)-ATR, followed by a significant decline to 4.5 mg/gfw after 14 days of exposure. The effect of Pb(II)-ATR was similar to that of Cd(II)-ATR but the amount of NP-SH was not higher than that of Cd(II)-ATR.

Tobacco nicotine uptake permease (NUP1) affects alkaloid metabolism.

An effective plant alkaloid chemical defense requires a variety of transport processes, but few alkaloid transporters have been characterized at the molecular level. Previously, a gene fragment encoding a putative plasma membrane proton symporter was isolated, because it was coordinately regulated with several nicotine biosynthetic genes. Here, we show that this gene fragment corresponds to a Nicotiana tabacum gene encoding a nicotine uptake permease (NUP1). NUP1 belongs to a plant-specific class of purine uptake permease-like transporters that originated after the bryophytes but before or within the lycophytes. NUP1 expressed in yeast cells preferentially transported nicotine relative to other pyridine alkaloids, tropane alkaloids, kinetin, and adenine. NUP1-GFP primarily localized to the plasma membrane of tobacco Bright Yellow-2 protoplasts. WT NUP1 transcripts accumulated to high levels in the roots, particularly in root tips. NUP1-RNAi hairy roots had reduced NUP1 mRNA accumulation levels, reduced total nicotine levels, and increased nicotine accumulation in the hairy root culture media. Regenerated NUP1-RNAi plants showed reduced foliar and root nicotine levels as well as increased seedling root elongation rates. Thus, NUP1 affected nicotine metabolism, localization, and root growth.

Soil penetration resistance, a suitable soil property to account for variations in root elongation and branching

Root system development is affected by soil conditions. The effects of bulk density, water content and penetration resistance on root development processes were investigated in peach trees. Peach tree rootstocks were grown in various soil conditions, combining two bulk densities (1.2 and 1.5 g soil.cm-3) and three water contents (0.14, 0.17 and 0.20 g.g-1soil). Root parameters (tip diameter, length of apical unbranched zone, branching density and diameters of main and lateral roots) and plant growth (leaves, branches, trunk, root dry mass) were measured. Root growth processes (elongation, branching) were studied using relationships between root parameters. The proportion of biomass allocated to each plant compartment was similar whatever the soil conditions. Variations in root development were best explained by the variation in penetration resistance, rather than other soil properties. Increased soil penetration resistance reduced the root elongation rate, especially for thick roots. In addition, the branching pattern was affected. In soil with a high penetration resistance, the root system shape differs from a typical herringbone pattern. These results allow quantification of the root system plasticity, and improve our understanding of the interactions between root development and soil properties.

A circadian and an ultradian rhythm are both evident in root growth of rice

This paper presents evidence for the existence of both a circadian and an ultradian rhythm in the elongation growth of rice roots. Root elongation of rice ( Oryza sativa) was recorded under dim green light by using a CCD camera connected to a computer. Four treatment conditions were set-up to investigate the existence of endogenous rhythms: 28 °C constant temperature and continuous dark (28 DD); 28 °C constant temperature and alternating light and dark (28 LD); 33 °C constant temperature and continuous dark (33 DD); and diurnal temperature change and alternating light and dark (DT-LD). The resulting spectral densities suggested the existence of periodicities of 20.4–25.2 h (circadian cycles) and 2.0–6.0 h (ultradian cycles) in each of the 4 treatments. The shorter ultradian cycles can be attributed to circumnutational growth of roots and/or to mucilage exudation. The average values across all the replicate data showed that the highest power spectral densities (PSDs) corresponded to root growth rhythms with periods of 22.9, 23.7, and 2.1 h for the 28 DD, 28 LD, and 33 DD treatments, respectively. Accumulation of PSD for each data set indicated that the periodicity was similar in both the 28 DD and 33 DD treatments. We conclude that a 23-h circadian and a 2-h ultradian rhythmicity exist in rice root elongation. Moreover, root elongation rates during the day were 1.08 and 1.44 times faster than those during the night for the 28 LD and DT-LD treatments, respectively.

Links between root developmental traits and foraging performance

We designed a simple dynamic and stochastic architectural model with six parameters to link the foraging performance of root systems to their developmental processes. Soil foraging was quantified by the volume enveloping the roots until a given uptake distance. Many simulated architectures were obtained by combining four different values for each parameter. The rate of soil colonization was mainly defined by individual root elongation rates and interbranch distances. Less intuitively, we showed that differentiation of elongation rates among the roots increased this colonization rate. Uptake efficiency--the ratio of the actual colonized volume to the volume of a unique cylinder with the same length and a radius corresponding to the uptake distance--declined with root system size. Nevertheless, large variations in efficiency existed among root systems for a given size, typically in a 4- to 10-fold range. Therefore, the 'efficiency gain' was defined as the deviation from the average trend in efficiency versus size. Between-root differentiation in elongation rates increased this gain. The level of hierarchy between mother and lateral roots, as well as the variation of elongation rates among lateral roots, was also shown to contribute to this optimization. Several parameter combinations could lead to similar efficiency gains.

Toxicity of metals to roots of cowpea in relation to their binding strength

Metal phytotoxicity is important in both environmental and agricultural systems. A solution culture study examined the toxicity of 26 metals to roots of cowpea (Vigna unguiculata (L.) Walp.); new data were collected for 15 metals and published data for 11 metals. Metal toxicity, calculated as causing a 50% reduction in root elongation rate, was determined based on either the measured concentration in the bulk solution (EC50(b)) or the calculated activity at the outer surface of the plasma membrane (EA50(0)°). The EC50(b) values ranged from 0.007 µM for Tl to 98,000 µM for K, with the order of rhizotoxicity to cowpea, from most to least toxic, being Tl = Ag > Cu > Hg = Ni = Ga = Ru = In > Sc = Cd = Gd = La = Co = Cs = Pb > Zn = Al = H > Mn > Ba = Sr > Li > Mg > Ca = Na > K. The EA50(0)° values suggest that the binding of metals to hard ligands is an important, general, nonspecific mechanism of toxicity, a hypothesis supported by the similar toxicity symptoms to roots of cowpea by many metals. However, additional mechanisms, such as strong binding to soft ligands, substantially increase rhizotoxicity of some metals, especially Tl, Ag, and Cs. Besides direct toxic effects, osmotic effects or reduced activity of Ca(2+) at the outer surface of the root plasma membrane (and resultant Ca deficiency) may decrease short-term root growth.

Calculated activity of Mn2+ at the outer surface of the root cell plasma membrane governs Mn nutrition of cowpea seedlings

Manganese (Mn) is an essential micronutrient for plant growth but is often toxic in acid or waterlogged soils. Using cowpea (Vigna unguiculata L. Walp.) grown with 0.05–1500 μM Mn in solution, two short-term (48 h) solution culture experiments examined if the effects of cations (Ca, Mg, Na, Al, or H) on Mn nutrition are related to the root cells’ plasma membrane (PM) surface potential, ψ00. When grown in solutions containing levels of Mn that were toxic, both relative root elongation rate (RRER) and root tissue Mn concentration were more closely related to the activity of Mn2+ at the outer surface of the PM, {Mn2+}00 (R2=0.812 and 0.871) than to its activity in the bulk solution, {Mn2+}b (R2=0.673 and 0.769). This was also evident at lower levels of Mn (0.05–10 μM) relevant to studies investigating Mn as an essential micronutrient (R2=0.791 versus 0.590). In addition, changes in the electrical driving force for ion transport across the PM influenced both RRER and the Mn concentration in roots. The {Mn2+}b causing a 50% reduction in root growth was found to be c. 500 to >1000 μM (depending upon solution composition), whilst the corresponding value was 3300 μM when related to {Mn2+}00. Although specific effects such as competition are not precluded, the data emphasize the importance of non-specific electrostatic effects in the Mn nutrition of cowpea seedlings over a 1×105-fold range of Mn concentration in solution.

Primary Root Elongation Rate and Abscisic Acid Levels of Maize in Response to Water Stress

Agricultural crops are exposed to greater water deficits as production regions receive less rainfall and producers move into more arid landscapes to meet demand. Maintenance of root elongation is vital for seedling establishment in these types of climates. Some maize (Zea mays L.) lines can maintain primary root elongation under severe water stress (SS) conditions. Twelve maize inbred lines were examined at the seedling stage before leaf expansion to assess the phenotypic diversity in primary root elongation rate and root elongation zone abscisic acid (ABA) content at three water potentials, −0.03 MPa, −0.3 MPa, and −1.6 MPa. A phylogenetic tree was constructed using 93 simple sequence repeat (SSR) markers to examine genetic and phenotypic relationships among the lines in relation to stress response. Statistical analysis revealed phenotypic diversity in both primary root elongation rate and ABA content. No correlations were detected between the amount of ABA present in the root tip and the rate of root elongation in any of the treatments. Histograms of primary root elongation response to the varying water deficits suggest multiple mechanisms may be responsible for the response to water stress observed in different maize lines. Different phenotypic responses toward water stress were observed for lines with small genetic distances, which will aid in identification of specific alleles for maintenance of primary root growth under water deficits.

Development of a novel semi-hydroponic phenotyping system for studying root architecture.

A semi-hydroponic bin system was developed to provide an efficient phenotyping platform for studying root architecture. The system was designed to accommodate a large number of plants in a small area for screening genotypes. It was constructed using inexpensive and easily obtained materials: 240L plastic mobile bins, clear acrylic panels covered with black calico cloth and a controlled watering system. A screening experiment for root traits of 20 wild genotypes of narrow-leafed lupin (Lupinus angustifolius L.) evaluated the reliability and efficiency of the system. Root architecture, root elongation rate and branching patterns were monitored for 6 weeks. Significant differences in both architectural and morphological traits were observed among tested genotypes, particularly for total root length, branch number, specific root length and branch density. Results demonstrated that the bin system was efficient in screening root traits in narrow-leafed lupin, allowing for rapid measurement of two-dimensional root architecture over time with minimal disturbance to plant growth and without destructive root sampling. The system permits mapping and digital measurement of dynamic growth of taproot and lateral roots. This phenotyping platform is a desirable tool for examining root architecture of deep root systems and large sets of plants in a relatively small space.

Root elongation, water stress, and mechanical impedance: a review of limiting stresses and beneficial root tip traits

Root elongation in drying soil is generally limited by a combination of mechanical impedance and water stress. Relationships between root elongation rate, water stress (matric potential), and mechanical impedance (penetration resistance) are reviewed, detailing the interactions between these closely related stresses. Root elongation is typically halved in repacked soils with penetrometer resistances >0.8-2 MPa, in the absence of water stress. Root elongation is halved by matric potentials drier than about -0.5 MPa in the absence of mechanical impedance. The likelihood of each stress limiting root elongation is discussed in relation to the soil strength characteristics of arable soils. A survey of 19 soils, with textures ranging from loamy sand to silty clay loam, found that ∼10% of penetration resistances were >2 MPa at a matric potential of -10 kPa, rising to nearly 50% >2 MPa at - 200 kPa. This suggests that mechanical impedance is often a major limitation to root elongation in these soils even under moderately wet conditions, and is important to consider in breeding programmes for drought-resistant crops. Root tip traits that may improve root penetration are considered with respect to overcoming the external (soil) and internal (cell wall) pressures resisting elongation. The potential role of root hairs in mechanically anchoring root tips is considered theoretically, and is judged particularly relevant to roots growing in biopores or from a loose seed bed into a compacted layer of soil.

Stable diurnal growth rhythms modulate root elongation of Arabidopsis thaliana

Arabidopsis root growth kinetics were investigated with high temporal and spatial resolution in combination with detailed statistical analysis to resolve presence of diurnal modulation of root tip displacement. In particular, high resolution video imaging was used to monitor root-tip displacement of Arabidopsis thaliana seedlings over several days. Root growth kinetics were sampled and statistically analyzed in two different photoperiods: long day LD (16 h-8 h) and equal LD (12 h-12 h) light. Diurnal root elongation kinetics exhibited five highly reproducible phases, one of these being a maximum of root growth rate displayed 1-2 h after the light on phase. Then, during the later part of the light period, root growth rate decreased. Several hours before darkening root elongation rates started to increase, with a profound decrease immediately after darkness. Subsequent to this dark-induced reduction in root growth rate the remaining part of the night was characterized by increasing growth activity. Together, tip elongation, in Arabidopsis roots is modulated by strong diurnal rhythms that are maintained in both photoperiods used and also in continuous illumination.

Short-term effects of coumarin along the maize primary root axis

The short-term effects of coumarin on three different maize primary root zones, transition zone (TZ, 3 mm), and two non-growing zones (NGZ1 and NGZ2 at 20 and 50 mm, respectively), were studied in order to investigate the effect of the allelochemical on maize root elongation rate (RER). The RER, plasma membrane (pm) H+-ATPase activity, quantitative pH changes and cell membrane potentials were evaluated. The results showed that coumarin caused at the TZ 1) an increased RER; 2) an enhancement of pm H+-ATPase activity and proton extrusion; and 3) a transient depolarization followed by a hyperpolarization of cell membrane potential. These observations were not evident in the NGZ1 and NGZ2 of the maize root. Coumarin-treatment in the NGZ1 did not change RER, but caused a membrane depolarization, while the NGZ2 was mostly insensitive to the allelochemical. These data suggested that the primary maize root was sensitive to coumarin within a 20 mm section from the root tip, but the more distal NGZ2 was not involved in coumarin-elicited physiological responses.

A potato NOA gene increased salinity tolerance in Arabidopsis thaliana

The purpose of this study is to produce recombinant StNOA1 in transgenic plants and to test its potential role in plant salt stress responses. The newly cloned StNOA1 gene from Solanum tuberosum L. was inserted into AtnOA1 mutant plant genome by Agrobaterium-mediated floral dip method. Transgene integration was verified by polymerase chain reaction (PCR) in 4 different lines of transgenic Atnoa1. Expression of StNOA1 gene was further analyzed by reverse trancription (RT)-PCR. Physiological analyses indicated that the transgenic line TL9 had higher proline, soluble protein and chlorophyll contents as well as lower content of malondialdehyde (MDA) compared to its receptor, Atnoa1 mutant, under salt stress condition. Root elongation and survival rate in TL9 were significantly higher than those in Atnoa1 seedlings under salt stress. Present study proved that StNOA1 participated in Arabidopsis thaliana salt stress responses and increased its salinity tolerance.

Rhizotoxic effects of silver in cowpea seedlings

Silver (Ag) is highly toxic to aquatic organisms, including algae, invertebrate animals, and fish, but little information exists on Ag rhizotoxicity in higher plants. In two solution culture experiments with approximately 1,000 microM Ca(NO3)2 and 5 microM H3BO3 (pH 5.4), 20 to 80% of added Ag (< or =2 microM) was lost from solution within approximately 30 min, with a further decrease after 48 h root growth. Using measured Ag concentrations at the start of the experiments, the median effective concentration (EC50) for root elongation rate of cowpea (Vigna unguiculata [L.] Walp. cv. Caloona) was 0.010 microM Ag in the first 4 h of exposure (0.021 microM in the first 8 h). This demonstrates that Ag (as Ag+) is rapidly rhizotoxic to cowpea seedlings at concentrations similar to those that are toxic to freshwater biota. Rupturing of rhizodermal and outer cortical layers was evident after 48 h with 0.13 to 0.57 microM Ag initially in solution, being most severe at 0.13 or 0.25 microM Ag. An additional experiment showed that ruptures were first evident after 20 h exposure to 0.17 microM Ag, with increased severity of rupturing over time. The rhizotoxic effects of Ag are similar to those of some other trace metals (e.g., Cu, Al, La) that bind strongly to hard ligands and weakly to soft ligands. The similarity of rupturing effects, despite the difference in strong binding to soft ligands by Ag and to hard ligands by the other metals, suggests a distinctive metabolic effect of Ag that binds only weakly to hard ligands.

Root growth dynamics of Aleppo pine (Pinus halepensis Mill.) seedlings in relation to shoot elongation, plant size and tissue nitrogen concentration

Large and high nitrogen (N) concentration seedlings frequently have higher survival and growth in Mediterranean forest plantations than seedlings with the opposite traits, which has been linked to the production of deeper and larger root systems in the former type of seedlings. This study assessed the influence of seedling size and N concentration on root growth dynamics and its relation to shoot elongation in Aleppo pine (Pinus halepensis Mill.) seedlings. We cultivated seedlings that differed in size and tissue N concentration that were subsequently transplanted into transparent methacrylate tubes in the field. The number of roots, root depth, and the root and shoot elongation rate (length increase per unit time) were periodically measured for 10 weeks. At the end of the study, we also measured the twig water potential (ψ) and the mass of plant organs. New root mass at the end of the study increased with seedling size, which was linked to the production of a greater number of new roots of lower specific length rather than to higher elongation rate of individual roots. Neither plant size nor N concentration affected root depth. New root mass per leaf mass unit, shoot elongation rate, and pre-dawn ψ were reduced with reduction in seedling size, while mid-day ψ and the root relative growth rate were not affected by seedling size. N concentration had an additive effect on plant size on root growth but its overall effect was less important than seedling size. Shoot and roots had an antagonistic elongation pattern through time in small seedlings, indicating that the growth of both organs depressed each other and that they competed for the same resources. Antagonism between shoot and root elongation decreased with plant size, disappearing in large and medium seedlings, and it was independent of seedling N concentration. We conclude that root and shoot growth but not rooting depth increased with plant size and tissue N concentration in Aleppo pine seedlings. Since production of new roots is critical for the establishment of planted seedlings, higher absolute root growth in large seedlings may increase their transplanting performance relative to small seedlings. The lack of antagonism between root and shoot growth in large seedlings suggests that these plants can provide resources to sustain simultaneous growth of both organs.

Analysis of Arabidopsis thaliana root growth kinetics with high temporal and spatial resolution

Methods exist to quantify the distribution of growth rate over the root axis. However, non-destructive, high-throughput evaluations of total root elongation in controlled environments and the field are lacking in growth studies. A new imaging approach to analyse total root elongation is described. High pixel resolution of the images enables the study of growth in short time intervals and provides high temporal resolution. Using the method described, total root elongation rates are calculated from the displacement of the root tip. Although the absolute root elongation rate changes in response to growth conditions, this set-up enables root growth of Arabidopsis wild-type seedlings to be followed for more than 1 month after germination. The method provides an easy approach to decipher root extension rate and much simpler calculations compared with other methods that use segmental growth to address this question. The high temporal resolution allows small modifications of total root elongation growth to be revealed. Furthermore, with the options to investigate growth of various mutants in diverse growth conditions the present tool allows modulations in root growth kinetics due to different biotic and abiotic stimuli to be unravelled. Measurements performed on Arabidopsis thaliana wild-type (Col0) plants revealed rhythms superimposed on root elongation. Results obtained from the starchless mutant pgm, however, present a clearly modified pattern. As expected, deviation is strongest during the dark period.

Survival of Schismus arabicus seedlings exposed to desiccation depends on annual periodicity

Schismus arabicus, a desert annual grass, is one of the most common pasture annuals in the deserts of Israel and Asia. S. arabicus exhibits a unique set of adaptations and survival strategies, which enable it to germinate, develop and produce seeds even in years with annual rainfall of less than 100 mm. The current study examined whether an annual rhythm exists in the survival ability of S. arabicus seedlings exposed to desiccation. Our results indicate that survival of S. arabicus seedlings after six different periods of 7 to 42 days of desiccation depended on the month of germination of the caryopses (seeds). Seed germination was 80-100% in all experiments, regardless the month of germination; however, seedlings that germinated in different months varied in their root and shoot elongation rates. None of about 2,500 seedlings that germinated in July (in each of the 4 years) survived the desiccation treatment. The percentages of surviving seedlings in each month of June from 2002 to 2005 were less than 40%. In contrast, over 80% of the seedlings that germinated in each of the months of December and January survived after the desiccation periods of 7-42 days. Seedlings that survived were transferred to 5 L soil pots in which the seedlings developed into mature plants, completed their life cycle and produced seeds that germinated well. The current study demonstrated a novel phenomenon indicating that seedling survival in plants may depend on an annual periodicity according to the date of germination.

Root Elongation Rate in Seedlings of Some Lines of Cowpea (Vigna unguiculata (L.) WALP.), Differing in Drought Tolerance

Root Elongation Rate in Seedlings of Some Lines of Cowpea (Vigna unguiculata (L.) WALP.), Differing in Drought Tolerance