An investigation of boron toxicity in barley using metabolomics

Boron (B) is an essential micronutrient that affects plant growth at either deficient or toxic concentrations in soil. The aim of this work was to investigate the adaptation of barley (Hordeum vulgare) plants to toxic B levels and to increase our understanding of B toxicity tolerance mechanisms. We used a metabolomics approach to compare metabolite profiles in root and leaf tissues of an intolerant, commercial cultivar (cv Clipper) and a B-tolerant Algerian landrace (cv Sahara). After exposure to elevated B (200 and 1,000 microM), the number and amplitude of metabolite changes in roots was greater in Clipper than in Sahara. In contrast, leaf metabolites of both cultivars only responded following 1,000 microM treatment, at which B toxicity symptoms (necrosis) were visible. In addition, metabolite levels were dramatically altered in the tips of leaves of the sensitive cultivar Clipper after growth in 1,000 microM B compared to those of Sahara. This correlates with a gradual accumulation of B from leaf base to tip in B-intolerant cultivars. Overall, there were always greater differences between tissue types (roots and leaves) than between the two cultivars. This work has provided insights into metabolic differences of two genetically distinct barley cultivars and information about how they respond metabolically to increasing B levels.     

Changes in hormonal balance and meristematic activity in primary root tips on the slowly rotating clinostat and their effect on the development of the rapeseed root system

The morphometry of the root system, the meristematic activity and the level of indole-3-acetic acid (IAA), abscisic acid (ABA) and zeatin in the primary root tips of rapeseed seedlings were analyzed as functions of time on a slowly rotating clinostat (1 rpm) or in the vertical controls (1 rpm). The fresh weight of the root system was 30% higher throughout the growth period (25 days) in clinorotated seedlings. Morphometric analysis showed that the increase in biomass on the clinostat was due to greater primary root growth, earlier initiation and greater elongation of the secondary roots, which could be observed even in 5-day-old seedlings. However, after 15 days, the growth of the primary root slowed on the clinostat, whereas secondary roots still grew faster in clinorotated plants than in the controls. At this time, the secondary roots began to be initiated closer to the root tip on the clinostat than in the control. Analysis of the meristematic activity and determination of the levels in IAA, ABA and zeatin in the primary root tips demonstrated that after 5 days on the clinostat, the increased length of the primary root could be the consequence of higher meristematic activity and coincided with an increase in both IAA and ABA concentrations. After 15 days on the clinostat, a marked increase in IAA, ABA and zeatin, which probably reached supraoptimal levels, seems to cause a progressive disturbance of the meristematic cells, inducing a decrease of primary root growth between 15 and 25 days. These modifications in the hormonal balance and the perturbation of the meristematic activity on the clinostat were followed by a loss of apical dominance, which was responsible for the early initiation of secondary roots, the greater elongation of the root system and the emergence of the lateral roots near the tip of the primary root.     

Phosphorus efficiencies and responses of barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L.) to arbuscular mycorrhizal fungi grown in highly calcareous soil

Two experiments were carried out to investigate phosphorus efficiencies and mycorrhizal responsiveness in an improved cultivar (Clipper) and a landrace (Sahara) of barley (Hordeum vulgare L.). In experiment 1, two pot sizes were used to evaluate the effect of soil volume on P uptake and mycorrhizal responsiveness. In experiment 2, a compartmented ("cross-pot") system was used to monitor (32)P delivery by external hyphae of arbuscular mycorrhizal fungi (AMF) to the host plant. Results showed that, irrespective of growth conditions, Sahara had much larger root biomass than Clipper and consequently substantially more P was allocated to roots in Sahara than in Clipper. Specific root length in Clipper was much longer than in Sahara. Increase in soil volume enhanced percentage root length colonised by AMF, plant growth and P uptake, and Sahara was more sensitive to changes in soil volume than Clipper. Pot size (soil volume) used to assess responsiveness to AMF by different plant species or genotypes with different root/shoot ratios might be a confounding factor. Clipper was more responsive to AMF than Sahara in terms of tissue P concentrations, which is partly related to their differences in root/shoot ratios. However, increases in SPU [specific P uptake, mg P (g root biomass)(-1)] caused by AMF were bigger in Clipper, suggesting that AMF played a larger role in P uptake. In accordance with the larger increase in SPU, Clipper took up more (32)P via AMF hyphae than Sahara. The compartmented system using radioactive P might be an alternative approach to directly investigate mycorrhizal responsiveness of different plant species or varieties than conventional pot experiments, provided that the same AM fungus is used.     

Genes mapping to boron tolerance QTL in barley identified by suppression subtractive hybridization

Boron tolerance is a quantitative trait controlled by multiple genes. Suppression subtractive hybridization was carried out on root cDNA from bulked boron tolerant and intolerant doubled haploid barley lines grown under moderate boron stress to identify genes associated with boron tolerance. One hundred and eleven clones representing known proteins were found to be up‐regulated in the tolerant bulk upon boron stress. Nine clones were genetically mapped to previously reported boron tolerance QTL. These include a clone identical to the boron transporter gene Bot1 and a clone coding for a bromo‐adjacent homology domain‐containing protein, mapping to the 6H boron tolerance locus and co‐segregating with reduced boron intake in a Clipper × Sahara‐3771 mapping population. A third clone mapping to the 2H QTL region encoding an S‐adenosylmethionine decarboxylase precursor was found to provide tolerance to high boron by heterologous expression. Yeast cells expressing Sahara SAMDC were able to grow on 15 mm boron solid media and maintained cellular boron concentrations at 13% lower than control cells expressing empty vector. The data suggest that an antioxidative response mechanism involving polyamines and the ascorbate–glutathione pathway in Sahara barley may provide an advantage in tolerating high soil concentrations of boron.     

Biochemical Changes Associated with the Growth of Root Tips of Vicia faba in vitro, and the Effect of 2,4-Dichlorophenoxyacetic Acid

Root tips of Vicia faba were cultured for 3 weeks on a semi-solidmedium containing 40 g/l sucrose ±10^–5 M 2:4-dichlorophenoxyaceticacid (2,4-D). Inhibition of root elongation and the stimulationof an actively dividing meristematic zone in the pericycle regionwere observed in the 2,4-D-treated tissues. Biochemical dataon the DNA content and respiration of these root tips correlatewell with the morphological observations. 2,4-D also induceda marked decrease in the -cellulose content of the cell wallsand analyses of the carbohydrate content of the ethanol-solublepool and starch content of the cultured root tips have providedtentative evidence for some of the controlling factors exertedin the presence and absence of 2,4-D.     

Effects of soil resistance to root penetration on leaf expansion in wheat (Triticum aestivum L.): kinematic analysis of leaf elongation

Wheat leaves (Triticum aestivum L.) elongated 50% more slowlywhen plants were grown in soils with high mechanical resistanceto penetration (R_s. The profiles of epidermal cell lengths alongthe growth zone of expanding leaves and the locations of newlyformed walls were recorded in order to compare the kineticsof elongation and partitioning of both meristematic and non-meristematiccells. In leaf 5, which completely developed under stress, highR_s, did not affect the flux of mature cells through the elongationzone; leaf elongation was reduced only because these cells wereshorter. This reduced size reflected a reduction in cell lengthat partitioning, associated with shorter cycling time. The relativerates of cell elongation before and after partitioning wereunchanged. Cell fluxes were similar because the population ofmeristematic cells was reduced, offsetting their increased partitioningrate. In contrast, in leaf 1, high R_s, had no effect on thenumber of dividing cells; elongation rate was reduced becauseof slower relative cell expansion rate and slower cell partitioningrate. These differences could reflect differences in the stageat which successive leaves perceived root stress and also time-dependentchanges in the responsiveness of leaf development to stress-inducedroot signals or in the nature of these signals. The data reveal that cell cycling time may in fact be decreasedby unfavourable growth conditions and is not directly relatedto cell expansion rates; they also show that the elongationrate of meristematic cells is partly independently controlledfrom that of non-meristematic cells. Key words: Wheat, kinematics of leaf expansion, cell partitioning, cell elongation, root impedance     

Boron alleviates aluminum toxicity in pea (Pisum sativum)

One important target of boron (B) deficiency and aluminum (Al) toxicity is cell wall. Thus we studied the hypothesis that B is capable of alleviating Al toxicity in pea (Pisum sativum). Short-term and prolonged Al exposure to pea roots at different B levels was carried out on uniform seedlings pre-cultured at a low B level. When seedlings with a low B level were supplied with or without B for 1 and 2 days before 24 h Al exposure, roots were longer while root diameter was thinner after B addition especially for 2 days even with exposure to Al; root elongation was inhibited while root diameter was enlarged by Al exposure. Callose induction by Al toxicity was higher with B added, but this was reversed after the removal of the cotyledons. Hematoxylin staining was lighter in the root tips given B, and Al content in the root tips and cell walls dropped after exposure to B. This indicates that B alleviated Al toxicity in the root tips during short-term Al exposure by decreasing Al binding in root cell walls. An increase in chlorophyll and biomass and reduced chlorosis were found at the higher level of B during prolonged Al treatment, which was coincided with the decreased Al contents, indicating that B alleviated Al toxicity to shoots. B supplementation alleviates some of the consequences of Al toxicity by limiting some Al binding in cell walls, resulting in less injury to the roots as well as less injury to the shoots.     

Effect of boron on cell elongation and division in squash roots

This work establishes that cessation of root elongation of intact squash (Cucurbita pepo L.) plants is an early result of boron deficiency. Root elongation is slowed by 6 hours and is virtually stopped as early as 24 hours after boron is first withheld from the nutrient solution. As root elongation ceased, cell elongation progressed distally into the region normally occupied by the apical meristem and eventually the meristem became indistinguishable. Differentiation was determined by use of an elongation index in which cell length was compared to cell width. This index ranged from a low of 0.8 in boron-sufficient root meristems to a high of 3 in root meristems grown in a boron-deficient nutrient solution for 98 hours. It is concluded that a continuous supply of boron is not essential for cell elongation but is required for maintenance of meristematic activity. Boron may act as a regulator of cell division in this tissue.     

Genetic and molecular analyses of resistance to a variant of Puccinia striiformis in barley

Seedlings of 62 Australian barley cultivars and two exotic barley genotypes were assessed for resistance to a variant of Puccinia striiformis, referred to as “Barley Grass Stripe Rust” (BGYR), first detected in Australia in 1998, which is capable of infecting wild Hordeum species and some genotypes of cultivated barley. Fifty-three out of 62 cultivated barley cultivars tested were resistant to the pathogen. Genetic analyses of seedling resistance to BGYR in six Australian barley cultivars and one Algerian barley landrace indicated that they carried either one or two major resistance genes to the pathogen. A single recessive seedling resistance gene, rpsSa3771, identified in Sahara 3771, was located on the long arm of chromosome 1 (7 H), flanked by the restriction fragment length polymorphism (RFLP) markers Xwg420 and Xcdo347 at genetic distances of 12.8 and 21.9 cM, respectively. Mapping resistance to BGYR at adult plant growth stages using the doubled haploid (DH) population Clipper × Sahara 3771 identified two major quantitative trait loci (QTL), one on the long arm of chromosome 3 (3 H) and the second on the long arm of chromosome 1 (7 H), accounting for 26 % and 18 % of the total phenotypic variation, respectively. The QTL located on chromosome 7HL corresponded to seedling resistance gene rpsSa3771 and the second QTL was concluded to correspond to a single APR gene, designated rpsCl, contributed by cultivar Clipper.     

Effect of Ethylene on Cell Division and Deoxyribonucleic Acid Synthesis in Pisum sativum

Ethylene and supraoptimal levels of 2,4-dichlorophenoxyacetic acid inhibit the growth of the apical hook region of etiolated Pisum sativum (var. Alaska) seedlings by stopping almost all cell divisions. Cells are prevented from entering prophase. The hormones also retard cell division in intact root tips and completely stop the process in lateral buds. The latter inhibition is reversed partially by benzyl adenine. In root tips and the stem plumular and subhook regions, ethylene inhibits DNA synthesis. The magnitude of this inhibition is correlated with the degree of repression of cell division in meristematic tissue, suggesting that the effect on cell division results from a lack of DNA synthesis. Ethylene inhibits cell division within a few hours with a dose-response curve similar to that for most other actions of the gas. Experiments with seedlings grown under hypobaric conditions suggest that the gas naturally controls plumular expansion and cell division in the apical region.     

Physiological response of plants to low boron

This review focuses on physiological responses in higher plants to B deficiency at the whole plant and organ level. Plants respond to decreasing B supply in soil solutions by slowing down or ceasing growth. Boron deficiency inhibits root elongation through limiting cell enlargement and cell division in the growing zone of root tips. In the case of severe B deficiency, the root cap, quiescent centre and protoderm of root tips disappear and root growth ceases, leading to the death of root tips. Although vascular bundles are weakly developed in B-deficient roots, early effects of B deficiency on their initiation and differentiation is poorly understood. Inhibited leaf expansion by low B indirectly decreases the photosynthetic capacity of plants, though exact roles of B in photosynthesis remain to be explored. The early inhibition of root growth, compared to shoot growth, increases the shoot:root ratio. It is hypothesised that this may enhance the susceptibility of plants to environmental stresses such as marginally deficient supplies of other nutrients and water deficit in soil.In the field, sexual reproduction is often more sensitive to low soil B than vegetative growth, and marked seed yield reductions can occur without symptoms being expressed during prior vegetative growth. In flowers, low B reduces male fertility primarily by impairing microsporogenesis and pollen tube growth. Post-fertilisation effects include impaired embryogenesis, resulting in seed abortion or the formation of incomplete or damaged embryos, and malformed fruit. However, there is a great diversity of effects of low B on reproductive growth among species, and within the same species between sites and seasons. Much of this diversity is not explained by the current literature. Key processes in reproductive development which may be impaired under B deficiency are proposed and discussed. These include the formation of a diverse array of cell wall types, the supply of carbohydrates for growth and storage reserves, and the production of flavonols. Inflorescence architecture, floral morphology, canopy structure and prevailing weather conditions are suggested as being important for xylem B delivery into flowers because of their impact on transpiration. The extent of phloem translocation of B into reproductive organs has yet to be fully assessed. The timing of B sensitive stages in reproduction of most crop plants need defining in order to facilitate appropriate timing of corrective B treatments.As most container studies have imposed B deficiency by withholding B, much of the data on severely B-deficient plants requires re-evaluation. Further studies are warranted to understand the effects of realistically low levels of B in solution on the growth of meristematic tissues and floral organs. A B-buffered solution culture system is recommended for some of this work.     

锌肥对不同基因型大麦吸收积累镉的影响

对土壤添加不同Zn、Cd条件下两种基因型(Sahara和Clipper)大麦对Zn、Cd的吸收积累研究表明,在本实验条件下土壤添加Zn、Cd对植物地上部生物量没有显著影响,但土壤添加Zn抑制植物根系生长,在土壤不缺Zn情况下添加Zn<20mg·kg^-1时并没有对大麦体内Cd浓度产生显著影响;当土壤Zn添加量达到40mg·kg^-1时,植物体内Cd浓度明显降低,植物吸收Cd的总量随着土壤添加Zn的增加而显著下降,这主要是由于根系生物量的下降所致,两个基因型大麦品种Zn效率存在显著差异,但这一差异对植物吸收Cd的总量没有影响,Zn高效品种Sahara根部Cd浓度显著低于Clipper。     

Ultrastructural detection of sucrose synthase distribution in developing maize leaves

Summary A developing maize leaf grows by the activity of a basal meristematic region and an adjacent elongating zone, resulting in a morphological and functional gradient along the leaf. We have used this system to detect the spatial and temporal expression of an enzyme, sucrose synthase, which plays a pivotal role in the sucrose import-export transition which occurs along a monocotyledon leaf. Immunogold labeling was used to detect the cellular and sub-cellular distribution of sucrose synthase (SS) at the electron microscopical level; the protein was visualized using a polyclonal antiserum on embedded tissue sections. Immunolabel was observed in the cytosol of dividing meristematic cells, expanding cells of the elongation zone, and in differentiating cells of young photosynthetic tissue. In fully differentiated leaf tissue, however, the protein was no longer immuno-detectable in photosynthetic cells, but was present in the guard and subsidiary cells of stomata and in companion cells within the phloem tissue of vascular bundles. The tissue- and cell-specific localization of sucrose synthase changes along the growing leaf as a function of the developmental state and the associated need for sucrose import or export.     

Gibberellic acid and dwarfism effects on the growth dynamics of B73 maize (Zea mays L.) leaf blades: a transient increase in apoplastic peroxidase activity precedes cessation of cell elongation

The relationship between apoplastic peroxidase (EC 1.11.1.7) activity and cessation of growth in maize (Zea mays L.) leaf blades was investigated by altering elongation zone length. Apoplastic peroxidase activity in the elongation and secondary cell wall deposition zones of elongating leaf blades of the maize inbred line B73 was used as a control and compared to leaves of the dwarf mutant D8-81127, a near-isogenic line of B73 unresponsive to gibberellins, and to leaves of B73 plants to which gibberellic acid (GA(3)) had been applied via root uptake. Elongation zone length was increased by treatment with GA(3) through an increase in cell number as well as increased final cell length. The shorter elongation zone of dwarf leaves occurred primarily through reduced final cell length. Although elongation zone length differed among dwarf, control, and GA(3)-treated leaf blades, in all three treatments a transient increase in apoplastic peroxidase activity preceded a reduction in the segmental elongation rate in leaves. A peroxidase isoenzyme with pI 7.0 occurred in the leaf elongation zone during growth deceleration in all three treatments, and its activity decreased as growth displaced tissue into the region of secondary cell wall deposition. Growth cessation for all treatments coincided with the first appearance of peroxidase isozymes with pIs of 5.6 and 5.7. Based on the activity of particular isozymes relative to growth and differentiation, the pI 7.0 isoenzyme is most likely to be involved in cessation of cell elongation, while isozymes with pIs 5.6 and 5.7 are likely to be active in lignification.     

Differential effects of sucrose and auxin on localized phosphate deficiency-induced modulation of different traits of root system architecture in Arabidopsis

Phosphorus, one of the essential elements for plants, is often a limiting nutrient in soils. Low phosphate (Pi) availability induces sugar-dependent systemic expression of genes and modulates the root system architecture (RSA). Here, we present the differential effects of sucrose (Suc) and auxin on the Pi deficiency responses of the primary and lateral roots of Arabidopsis (Arabidopsis thaliana). Inhibition of primary root growth and loss of meristematic activity were evident in seedlings grown under Pi deficiency with or without Suc. Although auxin supplementation also inhibited primary root growth, loss of meristematic activity was observed specifically under Pi deficiency with or without Suc. The results suggested that Suc and auxin do not influence the mechanism involved in localized Pi sensing that regulates growth of the primary root and therefore delineates it from sugar-dependent systemic Pi starvation responses. However, the interaction between Pi and Suc was evident on the development of the lateral roots and root hairs in the seedlings grown under varying levels of Pi and Suc. Although the Pi+ Suc- condition suppressed lateral root development, induction of few laterals under the Pi- Suc- condition point to increased sensitivity of the roots to auxin during Pi deprivation. This was supported by expression analyses of DR5uidA, root basipetal transport assay of auxin, and RSA of the pgp19 mutant exhibiting reduced auxin transport. A significant increase in the number of lateral roots under the Pi- Suc- condition in the chalcone synthase mutant (tt4-2) indicated a potential role for flavonoids in auxin-mediated Pi deficiency-induced modulation of RSA. The study thus demonstrated differential roles of Suc and auxin in the developmental responses of ontogenetically distinct root traits during Pi deprivation. In addition, lack of cross talk between local and systemic Pi sensing as revealed by the seedlings grown under either the Pi- Suc- condition or in the heterogeneous Pi environment highlighted the coexistence of Suc-independent and Suc-dependent regulatory mechanisms that constitute Pi starvation responses.     

Inhibition of maize root growth by high nitrate supply is correlated with reduced IAA levels in roots

The plant root system is highly sensitive to nutrient availability and distribution in the soil. For instance, root elongation is inhibited when grown in high nitrate concentrations. To decipher the mechanism underlying the nitrate-induced inhibition of root elongation, the involvement of the plant hormone auxin in nitrate-dependent root elongation of maize was investigated. Root growth, nitrogen and nitrate concentrations, and indole-3-acetic acid (IAA) concentrations in roots and in phloem exudates of maize grown under varying nitrate concentrations were analyzed. Total N and nitrate concentrations in shoots and roots increased and elongation of primary, seminal and crown roots were inhibited with increasing external nitrate from 0.05 to 5 mM. High nitrate-inhibited root growth resulted primarily from the reduced cell elongation and not from changes in meristem length. IAA concentrations in phloem exudates reduced with higher nitrate supply. Inhibition of root growth by high nitrate was closely related to the reduction of IAA levels in roots, especially in the sections close to root tips. Exogenous NAA and IAA restored primary root growth in high nitrate concentrations. It is concluded that the inhibitory effect of high nitrate concentrations on root growth may be partly attributed to the decrease in auxin concentrations of roots.     

Specific Leaf Weight in Zones of Cell Division, Elongation and Maturation in Tall Fescue Leaf Blades

Patterns of change in specific leaf weight (SLW), water-solublecarbohydrate (WSC) content and leaf width were used to delineatethe region of secondary cell wall accumulation, and determinethe rate of increase in structural material along a developingleaf blade of tall fescue (Festuca arundinacea Schreb.). Structuralspecific leaf weight (SSLW) was determined by subtracting WSCmass from dry weight to emphasize structural material. Becausemeristematic activity, cell elongation, and cellular maturationare arranged successively in the grass leaf, these patternsrepresent a developmental sequence through which each segmentof the leaf blade passes. Patterns were generally similar fortwo genotypes, one selected for high (HYT) and the other forlow (LYT) yield per tiller, for a single genotype grown at 17or 25 C, and for two field-grown populations which differedin leaf area expansion rate (LAER). In all three studies, the elongation zone of the developingleaf had 31 to 39 per cent WSC on a dry weight basis. The LYTgenotype had a higher SLW at all stages of development whengrown at 17 than at 25 C, due to greater WSC accumulation.At 20 C, the HYT genotype had a higher SLW all along the elongatingleaf blade than the LYT genotype. This difference was due toa difference in SSLW, while WSC content was similar. The LERwas 64 per cent higher in the high population than the low,but elongation zones were similar in WSC. In all cases, SSLWwas high in the meristematic region, lowest near the distalend of the cell elongation zone, then increased linearly astissue matured. Rate of increase in SSLW was 8.5 and 5.2 g m^–2d^–1 for the HYT and LYT genotypes, respectively, and 7.6and 6.7 g m^–2 d^–1 for the high and low LAER populations,respectively. Festuca arundinacea Schreb., tall fescue, specific leaf weight, leaf width, water-soluble carbohydrates, leaf elongation rate     

Hardening of root cell walls: a growth inhibitory response to salinity stress

Primary roots of intact maize plants (Zea mays L.) grown for several days in nutrient solutions containing 100 mol m⁻³ NaCl and additional calcium, had relatively inhibited rates of elongation. Possible physical restraints underlying this salt induced inhibition were investigated. The inhibition did not involve reductions in osmotic potential gradients and turgor in the tip tissues responsible for root elongation growth. The apparent yield threshold pressure, which is related to capacity of cell walls to undergo loosening by stress relaxation, was estimated psychrometrically in excised root tips. Salinity increased yield threshold values. Comparative root extensibility values were obtained for intact plants by determining the initial (1 min) increase in root elongation rate induced by an 0.1 MPa osmotic jump. Comparative extensibility was significantly reduced in the salinized root tips. Salinity did not reduce capacities for water efflux and associated elastic contraction in root tip tissues of intact plants exposed to hypertonic mannitol. We conclude that cell wall hardening in the elongating root tips is an important component of root growth inhibition induced by long-term salinization.     

Mapping of chromosome regions conferring boron toxicity tolerance in barley (Hordeum vulgare L.)

 Boron toxicity has been recognised as an important problem limiting production in the low-rainfall regions of southern Australia, West Asia and North Africa. Genetic variation for boron toxicity tolerance in barley has been characterised but the mode of inheritance and the location of genes controlling tolerance were not previously known. A population of 150 doubled-haploid lines from a cross between a boron toxicity tolerant Algerian landrace, Sahara 3771, and the intolerant Australian cultivar Clipper was screened in four tolerance assays. An RFLP linkage map of the Clipper×Sahara population was used to identify chromosomal regions associated with boron tolerance in barley. Interval regression-mapping allowed the detection of four chromosomal regions involved in the boron tolerance traits measured. A region on chromosome 2H was associated with leaf-symptom expression, a region on chromosome 3H was associated with a reduction of the affect of boron toxicity on root growth suppression, a region on chromosome 6H was associated with reduced boron uptake, and a region on chromosome 4H was also associated with the control of boron uptake as well as being associated with root-length response, dry matter production and symptom expression. The benefits and potential of marker-assisted selection for boron toxicity tolerance are discussed. Received: 18 December 1997 / Accepted: 28 November 1998