Mechanical stimulation in Brachypodium distachyon: Implications for fitness,productivity, and cell wall properties

Mechanical stimulation, including exposure to wind, is a common environmental variable for plants. However, knowledge about the morphogenetic response of the grasses (Poaceae) to mechanical stimulation and impact on relevant agronomic traits is very limited. Two natural accessions of Brachypodium distachyon were exposed to wind and mechanical treatments. We surveyed a wide range of stem-related traits to determine the effect of the two treatments on plant growth, development, and stem biomass properties. Both treatments induced significant quantitative changes across multiple scales, from the whole plant down to cellular level. The two treatments resulted in shorter stems, reduced biomass, increased tissue rigidity, delayed flowering, and reduced seed yield in both accessions. Among changes in cell wall-related features, a substantial increase in lignin content and pectin methylesterase activity was most notable. Mechanical stimulation also reduced the enzymatic sugar release from the cell wall, thus increasing biomass recalcitrance. Notably, treatments had a distinct and opposite effect on vascular bundle area in the two accessions, suggesting genetic variation in modulating these responses to mechanical stimulation. Our findings highlight that exposure of grasses to mechanical stimulation is a relevant environmental factor affecting multiple traits important for their utilization in food, feed, and bioenergy applications.     

Shifts in fine root traits within and among species along a fine-scale hydrological gradient

Background and AimsLessons from above-ground trait ecology and resource economics theory may not be directly translatable to below-ground traits due to differences in function, trade-offs and environmental constraints. Here we examine root functional traits within and across species along a fine-scale hydrological gradient. We ask two related questions: (1) What is the relative magnitude of trait variation across the gradient for within- versus among-species variation? (2) Do correlations among below-ground plant traits conform with predictions from resource-economic spectrum theory?MethodsWe sampled four below-ground fine-root traits (specific root length, branching intensity, root tissue density and root dry matter content) and four above-ground traits (specific leaf area, leaf size, plant height and leaf dry matter content) in vascular plants along a fine-scale hydrological gradient within a wet heathland community in south-eastern Australia. Below-ground and above-ground traits were sampled both within and among species.Key ResultsRoot traits shifted both within and among species across the hydrological gradient. Within- and among-species patterns for root tissue density showed similar declines towards the wetter end of the gradient. Other root traits showed a variety of patterns with respect to within- and among-species variation. Filtering of species has a stronger effect compared with the average within-species shift: the slopes of the relationships between soil moisture and traits were steeper across species than slopes of within species. Between species, below-ground traits were only weakly linked to each other and to above-ground traits, but these weak links did in some cases correspond with predictions from economic theory.ConclusionsOne of the challenges of research on root traits has been considerable intraspecific variation. Here we show that part of intraspecific root trait variation is structured by a fine-scale hydrological gradient, and that the variation aligns with among-species trends in some cases. Patterns in root tissue density are especially intriguing and may play an important role in species and individual response to moisture conditions. Given the importance of roots in the uptake of resources, and in carbon and nutrient turnover, it is vital that we establish patterns of root trait variation across environmental gradients.     

Diallelic analysis of quantitative traits in hexaploid wheat (Triticum aestivum L.)

Abstract

A complete diallel study of crosses between eight wheat varieties was carried out to determine the relative magnitude of components of genetic variation and heritability for important grain yield, quality and drought‐related traits. The data appeared adequate for the additive‐dominance model. The additive effects predominated for most traits, and consequently the narrow‐sense heritability was high to moderately high for flag leaf area, weight and venation, stomatal frequency and size, epidermal cell size, biomass, protein content, number of tillers, spike length, spike density, 1000‐grain weight and grain yield. These results appear promising for selecting better plants in the segregating populations with some degree of improvement for yield, quality and physiological efficiency.     

Mechanical Differences Between Free-standing and Supported Wheat Plants, Triticum aestivum L.

The effect of wind sway on the mechanical characteristics ofthe anchorage roots and the stem was investigated in maturewinter wheat (Triticum aestivumL., cv. Hereward). Wheat plantswere field-grown, either supported by a frame, which preventedwind sway, or unsupported (free-standing) and the morphologyand mechanical properties of the stems and the anchorage, ‘coronal’, roots were measured. Wind sway had little influence on either the stem height orear weight of the plants but did affect the mechanical propertiesof the stem. Stems of supported plants were weaker and moreflexible than the stems of free-standing plants. There werealso differences in the anchorage systems between the treatments:supported plants had just under half as many ‘coronal’ anchorage roots as the free-standing plants. This reducedthe anchorage strength of supported plants by a third. These differences in mechanical structure meant that the free-standingplants were more resistant to stem buckling and more resistantto anchorage failure. However, considering the difference inthe need for mechanical strength in plants from the two regimes,these differences were small. This suggests that wheat has inherentmechanical integrity and, as a monocotyledon with no secondarythickening, it differs little structurally between environments. Triticum aestivumL.; thigmomorphogenesis; anchorage; safety factor; mechanical stimulation     

The influence of light conditions and interspecific competition on the root foraging traits and seedling growth of two tree species

Abstract

Enriched nutrient patches within natural soil represent an important source of nutrients for tree growth. In the present study, pot experiments in a heterogeneous nutrient environment were conducted to investigate the influence of light conditions and interspecific competition on the root foraging traits and seedling growth of Pinus massoniana and Schima superba. The root foraging scale and the whole-seedling biomass of both species were decreased by shading. The result of this treatment was a lower sensitivity to nutrient heterogeneity in plants that underwent the shading treatment than in plants that were exposed to full-light conditions. The above-ground biomass and whole-seedling biomass of S. superba were not affected by competition with P. massoniana. In contrast, the above-ground biomass and whole-seedling biomass of P. massoniana were negatively affected by competition with S. superba. The more rapid rate of root extension and the more efficient resource uptake of S.superba appear to explain this effect. The species-specific patterns of the influence of environmental factors on foraging ability and seedling growth should be given thorough consideration and should be applied to afforestation and to the management of tree plantations.     

Interactive effects of mechanical stress,sand burial and defoliation on growth and mechanical properties in Cynanchum komarovii

In drylands, wind, sand burial and grazing are three important factors affecting growth and mechanical properties of plants, but their interactive effects have not yet been investigated. Plants of the semi‐shrub Cynanchum komarovii, common in semi‐arid parts of NE Asia, were subjected to brushing, burial and defoliation. We measured biomass allocation and relative increment rates of dry mass (RGR_m), height (RGR_h) and basal diameter (RGR_d). We also measured the stem mechanical properties, Young’s modulus (E), second moment of area (I), flexural stiffness (EI) and breaking stress (σ_b), and scaled these traits to the whole‐plant level to determine the maximum lateral force (F_lateral) and the buckling safety factor (BSF). Brushing increased RGR_m; neither burial nor defoliation independently affected RGR_m, but together they reduced it. Among buried plants, brushing positively affected stem rigidity and strength through increasing RGR_d, E, I and EI, and at whole plant level this resulted in a larger BSF and F_lateral. However, among unburied plants this pattern was not observed. Our results thus show that effects of mechanical stress and grazing on plants can be strongly modified by burial, and these interactions should be taken into account when considering adaptive significance of plant mechanical traits in drylands.     

Relationships between carbon isotope discrimination, dry matter production, and harvest index in durum wheat

Carbon isotope discrimination has been proposed as a criterion for the indirect selection to improve transpiration efficiency and grain yield in bread wheat and barley. Less attention has been devoted to durumwheat (Triticum durumDesf.) despite its economic importance in the Mediterranean basin. The Δ genetic variation and its relationships to dry matter production and harvest index in durum wheat were investigated in this study. For this purpose, field experiments were conducted on 144 durum wheat accessions under Mediterranean conditions (South of France) during three consecutive years with contrasting climatic conditions. Grain yield, above-ground biomass, harvest index, and carbon isotope discrimination of flag leaf and kernel were measured. Differences between years, noted for both leaf and kernel carbon isotope discrimination, were probably related to the variation in water availability from year to year. A large genotypic variation was also noticed for both leaf and kernel carbon isotope discrimination. The two traits were found to be positively correlated with grain yield within and across years, which confirms the interest in carbon isotope discrimination for selection for grain yield improvement under Mediterranean conditions. Both kernel and leaf carbon isotope discrimination correlated better with harvest index than with grain yield, suggesting that carbon isotope discrimination could reflect the efficiency of carbon partitioning to the kernel. The lack of correlation between leaf carbon isotope discrimination and both harvest index and grain yield in favourable water conditions (1996) was probably due to the difference in water availability between the period until flag leaves sampling (favourable conditions) and the strong water stress which accompanied the grain filling. Kernel carbon isotope discrimination correlated better with both harvest index and grain yield than did leaf carbon isotope discrimination. Moreover, a higher broad-sense heritability was obtained for kernel carbon isotope discrimination than for leaf carbon isotope discrimination. As a result, kernel carbon isotope discrimination appeared to be a better predictive criterion for efficiency of the carbon partitioning to the kernel (harvest index), and hence for grain yield, than did flag leaf carbon isotope discrimination.     

Variation in phosphorus and sulfur content shapes the genetic architecture and phenotypic associations within the wheat grain ionome

Dissection of the genetic basis of wheat ionome is crucial for understanding the physiological and biochemical processes underlying mineral accumulation in seeds, as well as for efficient crop breeding. Most of the elements essential for plants are metals stored in seeds as chelate complexes with phytic acid or sulfur‐containing compounds. We assume that the involvement of phosphorus and sulfur in metal chelation is the reason for strong phenotypic correlations within ionome. Adjustment of element concentrations for the effect of variation in phosphorus and sulfur seed content resulted in drastic change of phenotypic correlations between the elements. The genetic architecture of wheat grain ionome was characterized by quantitative trait loci (QTL) analysis using a cross between durum and wild emmer wheat. QTL analysis of the adjusted traits and two‐trait analysis of the initial traits paired with either P or S considerably improved QTL detection power and accuracy, resulting in the identification of 105 QTLs and 617 QTL effects for 11 elements. Candidate gene search revealed some potential functional associations between QTLs and corresponding genes within their intervals. Thus, we have shown that accounting for variation in P and S is crucial for understanding of the physiological and genetic regulation of mineral composition of wheat grain ionome and can be implemented for other plants.     

Brushing effects on the growth and mechanical properties of Corispermum mongolicum vary with water regime

High water availability and mechanical stress can induce opposite responses in plants. In arid areas of Northern China the occurrence of high wind and high water availability tend to be negatively correlated. Since turgor pressure is a determinant of the mechanical stability of annuals, it is hypothesised that the effects of mechanical perturbation (MP) on annuals may depend on soil water availability. To test this proposal, we conducted an experiment in which a pioneering annual Corispermum mongolicum was subjected to two levels of MP and water supply, and then determined its growth and mechanical traits. Brushing had no effect on plant height and total biomass, but stimulated leaf and branch production. Water supply affected plant height, basal diameter, total biomass and stem rigidity, but not leaf and branch number, root/shoot ratio or flexibility. With high water availability, brushing stimulated the production of stiffer stems (thicker and with a higher Young's modulus) and more roots relative to shoot mass, but with low water availability MP induced the opposite response. This shows that both the degree and direction of plant responses to MP depend on the presence of other factors. We discuss how the interactive effects of MP and water availability on growth and mechanical properties may help C. mongolicum to establish in windy and arid environments.     

Plasticity to wind is modular and genetically variable in Arabidopsis thaliana

Thigmomorphogenesis, the characteristic phenotypic changes by which plants react to mechanical stress, is a widespread and probably adaptive type of phenotypic plasticity. However, little is known about its genetic basis and population variation. Here, we examine genetic variation for thigmomorphogenesis within and among natural populations of the model system Arabidopsis thaliana. Offspring from 17 field-collected European populations was subjected to three levels of mechanical stress exerted by wind. Overall, plants were remarkably tolerant to mechanical stress. Even high wind speed did not significantly alter the correlation structure among phenotypic traits. However, wind significantly affected plant growth and phenology, and there was genetic variation for some aspects of plasticity to wind among A. thaliana populations. Our most interesting finding was that phenotypic traits were organized into three distinct and to a large degree statistically independent covariance modules associated with plant size, phenology, and growth form, respectively. These phenotypic modules differed in their responsiveness to wind, in the degree of genetic variability for plasticity, and in the extent to which plasticity affected fitness. It is likely, therefore, that thigmomorphogenesis in this species evolves quasi-independently in different phenotypic modules.     

Yield vs. Quality trade‐offs for wheat in response to carbon dioxide and ozone

Although it is established that there exist potential trade‐offs between grain yield and grain quality in wheat exposed to elevated carbon dioxide (CO₂) and ozone (O₃), their underlying causes remain poorly explored. To investigate the processes affecting grain quality under altered CO₂ and O₃, we analysed 57 experiments with CO₂ or O₃ exposure in different exposure systems. The study covered 24 cultivars studied in 112 experimental treatments from 11 countries. A significant growth dilution effect on grain protein was found: a change in grain yield of 10% by O₃ was associated with a change in grain protein yield of 8.1% (R² = 0.96), whereas a change in yield effect of 10% by CO₂ was linked to a change in grain protein yield effect of 7.5% (R² = 0.74). Superimposed on this effect, elevated CO₂, but not O₃, had a significant negative effect on grain protein yield also in the absence of effects on grain yield, indicating that there exists a process by which CO₂ restricts grain protein accumulation, which is absent for O₃. Grain mass, another quality trait, was more strongly affected by O₃ than grain number, whereas the opposite was true for CO₂. Harvest index was strongly and negatively influenced by O₃, but was unaffected by CO₂. We conclude that yield vs. protein trade‐offs for wheat in response to CO₂ and O₃ are constrained by close relationships between effects on grain biomass and less than proportional effects on grain protein. An important and novel finding was that elevated CO₂ has a direct negative effect on grain protein accumulation independent of the yield effect, supporting recent evidence of CO₂‐induced impairment of nitrate uptake/assimilation. Finally, our results demonstrated that processes underlying responses of grain yield vs. quality trade‐offs are very different in wheat exposed to elevated O₃ compared with elevated CO₂.     

Effect of mechanical perturbation on the biomechanics, primary growth and secondary tissue development of inflorescence stems of Arabidopsis thaliana

Background and Aims

Mechanical perturbation is known to inhibit elongation of the inflorescence stem of Arabidopsis thaliana. The phenomenon has been reported widely for both herbaceous and woody plants, and has implications for how plants adjust their size and form to survive in mechanically perturbed environments. While this response is an important aspect of the plant''s architecture, little is known about how mechanical properties of the inflorescence stem are modified or how its primary and secondary tissues respond to mechanical perturbation.

Methods

Plants of the Columbia-0 ecotype were exposed to controlled brushing treatments and then submitted to three-point bending tests to determine stem rigidity and stiffness. Contributions of different tissues to the inflorescence stem geometry were analysed.

Key Results

Perturbed plants showed little difference in stem diameter, were 50 % shorter, 75 % less rigid and 70 % less stiff than controls. Changes in mechanical properties were linked to significant changes in tissue geometry – size and position of the pith, lignified interfascicular tissue and cortex – as well as a reduction in density of lignified cells. Stem mechanical properties were modified by changes in primary tissues and thus differ from changes observed in most woody plants tested with indeterminate growth – even though a vascular cambium is present in the inflorescence axis.

Conclusions

The study suggests that delayed development of key primary developmental features of the stem in this ecotype of Arabidopsis results in a ‘short and flexible’ rather than a ‘short and rigid’ strategy for maintaining upright axes in conditions of severe mechanical perturbation. The mechanism is comparable with more general phenomena in plants where changes in developmental rate can significantly affect the overall growth form of the plant in both ecological and evolutionary contexts.     

Grain set failure in boron deficient wheat

Effects of boron (B) deficiency on reproductive development and grain set in wheat was studied in experiments in a sand culture in which grain set was increased by increasing B supply in the nutrient solution. Early vegetative response was also studied in a solution culture experiment with 5 M B and without added B. Effects of B deficiency on the male and female part of the wheat flower were studied in a cross fertilization experiment involving B deficient and B sufficient wheat plants. An international trial (the Boron Probe Nursery) was conducted as a collaboration between Chiang Mai University, CIMMYT and National Agricultural Research Systems in various countries, to verify the B response in non-traditional, warm wheat-growing areas.There was a wide genotypic variation in reproductive responses to B among the eight wheat genotypes studied. In sand culture with low B (0.2 M), grain set index ranged from 9.5% in SW41 to 94.5% in Fang 60; with high B (10 M) it was 90% in all genotypes. Early vegetative response to B was measured in the length of the youngest emerged blade at 12 days after sowing. Without added B the length of the leaf blade relative to that with 5 M B ranged from 0.82 to 0.92. This indicates some variation in vegetative response to B among the genotpes. However, there was no relationship between vegetative and reproductive responses to B of the wheat genotypes.Fertility of both the male and female part of the wheat flower appears to be affected by B deficiency. Ears from B deficient plants that were bagged to prevent cross fertilization set no grain. Cross pollination of B deficient female flowers with pollen from B sufficient plants resulted in only 28% grain set, compared with 94% percent from manual crossing of B sufficient pollen on B sufficient female.Reponses to B application of SW41 and other sensitive genotypes at field sites of the first international Boron Probe Nursery (1990/91) confirmed that B deficiency can be a major cause of grain set failure in wheat in warm areas.     

Inheritance of the traits of spike and grain color and morphometric parameters of the grain in F<Subscript>1</Subscript> and F<Subscript>2</Subscript> hybrids of <Emphasis Type="Italic">Triticum spelta</Emphasis> L. × <Emphasis Type="Italic">Triticum aestivum</Emphasis> L.

The character of inheritance of the morphological traits of spike and grain color and morphometric parameters of the grain in simple and backcross F₁ and F₂ hybrids of spelt and soft wheat has been investigated. The experiments confirmed that single homologous genes determine the trait of grain width in different species of wheat. Incomplete dominance of the gene that determines the trait of grain length has been revealed. The increase of the dosage of genes from one wheat species in a backcross hybrid has been shown to increase the deviation from the other species and to bring the values of the quantitative parameters of the grain closer to the values for the saturating species. Splitting of the spike color trait in the F₂ offspring has been shown to follow the 15: 1 dihybrid cross scheme and to be controlled by two dominant homologous genes.     

The Effects of Selenium on Physiological Traits, Grain Selenium Content and Yield of Winter Wheat at Different Development Stages

The paper evaluated the effects of Se application time and rate on physiological traits, grain Se content, and yield of winter wheat by field experiment. Se application significantly increased grain Se content and yield, and the increased amount treated with 20 and 30 mg Se?L^?1 was the highest. At blooming–filling stage, Se application significantly increased grain Se content, but did not affect yield. Chlorophyll content was increased by Se application, and the increased amount at heading–blooming stage was higher than that in wheat leaves at the other stages. At four development stages, Se treatments (except for 10 mg Se?L^?1 at jointing–heading stage) significantly decreased the rate of superoxide (O₂ ^?) radical production. At heading–blooming (except for 50 mg Se?L^?1) and blooming–filling stages, hydrogen peroxide (H₂O₂) content was significantly decreased by Se treatments. The rate of O₂ ^? production and H₂O₂ content at 20 and 30 mg Se?L^?1 was the lowest. Se treatments (except for 10 mg Se?L^?1 at regreening–jointing and blooming–filling stages) also induced an evident decrease in malondialdehyde content. Proline content induced by Se treatments at jointing–heading and heading–blooming stages was higher than that in wheat leaves at regreening–jointing and blooming–filling stages. At four development stages, Se treatments all significantly increased glutathione peroxidase activity, and the treatments with 20 and 30 mg Se?L^?1 also evidently increased reduced glutathione content. These results suggested that Se application at different development stages increased antioxidant capacity of wheat, reduced oxidant stress to some extent, and the effects of Se treatments was the best if Se concentration ranged between 20 and 30 mg Se?L^?1. In addition, Se application time was more beneficial for Se accumulation and yield in wheat grain at heading–blooming stage.     

Increased chilling tolerance and altered carbon metabolism in tomato leaves following application of mechanical stress

We investigated the effects of brushing on the chilling tolerance and metabolism of nonstructural carbohydrates (soluble sugars and starch) in tomato leaves before, during and after a chilling stress. Tomato plants ( Lycopersicon esculentum Mill. cv. Caruso) were cultivated either without mechanical stress application (control plants) or with daily brushing treatments for 15 days (brushed plants), prior to a 7-day chilling treatment (8/5°C day/night). Brushing resulted in shorter plants with a 34% reduction in leaf dry weight per area and a 59% reduction of soluble sugars and starch, on a dry weight basis. The sugar to starch ratio was not affected by brushing. A greater chilling tolerance in the brushed plants was demonstrated by the maintenance of a significantly higher PSII efficiency in brushed plants (42%) compared to that of the control plants (30%) after 7 days of chilling treatment, less visible damage to the leaf tissue, and a more rapid resumption of growth during 3 days of recovery as compared to control plants. During the chilling treatment levels of soluble sugars per leaf dry weight increased 15-fold in the brushed plants and 5-fold in control plants. In the present study we have demonstrated that brushing can increase chilling tolerance in tomato plants. The observed differences in chilling tolerance and concentration of soluble sugars in the leaves may indicate an involvement of soluble sugar levels in acclimation to chilling.     

Direct and indirect selection on flowering time,water‐use efficiency (WUE, δ13C), and WUE plasticity to drought in Arabidopsis thaliana

Flowering time and water-use efficiency (WUE) are two ecological traits that are important for plant drought response. To understand the evolutionary significance of natural genetic variation in flowering time, WUE, and WUE plasticity to drought in Arabidopsis thaliana, we addressed the following questions: (1) How are ecophysiological traits genetically correlated within and between different soil moisture environments? (2) Does terminal drought select for early flowering and drought escape? (3) Is WUE plasticity to drought adaptive and/or costly? We measured a suite of ecophysiological and reproductive traits on 234 spring flowering accessions of A. thaliana grown in well-watered and season-ending soil drying treatments, and quantified patterns of genetic variation, correlation, and selection within each treatment. WUE and flowering time were consistently positively genetically correlated. WUE was correlated with WUE plasticity, but the direction changed between treatments. Selection generally favored early flowering and low WUE, with drought favoring earlier flowering significantly more than well-watered conditions. Selection for lower WUE was marginally stronger under drought. There were no net fitness costs of WUE plasticity. WUE plasticity (per se) was globally neutral, but locally favored under drought. Strong genetic correlation between WUE and flowering time may facilitate the evolution of drought escape, or constrain independent evolution of these traits. Terminal drought favored drought escape in these spring flowering accessions of A. thaliana. WUE plasticity may be favored over completely fixed development in environments with periodic drought.