Cessation of tillering in spring wheat in relation to light interception and red : far-red ratio

BACKGROUND AND AIMS: The production of axillary shoots (tillering) in spring wheat (Triticum aestivum) depends on intraspecific competition. The mechanisms that underlie this competition are complex, but light within the wheat canopy plays a key role. The main objectives of this paper are to analyse the effects of plant population density and shade on tillering dynamics of spring wheat, to assess the canopy conditions quantitatively at the time of tillering cessation, and to analyse the relationship between the tiller bud and the leaf on the same phytomer. METHODS: Spring wheat plants were grown at three plant population densities and under two light regimes (25 % and 100 % light). Tiller appearance, fraction of the light intercepted, and red : far-red ratio at soil level were recorded. On six sampling dates the growth status of axillary buds was analysed. KEY RESULTS: Tillering ceased earlier at high population densities and ceased earlier in the shade than in full sunlight. At cessation of tillering, both the fraction of light intercepted and the red : far-red ratio at soil level were similar in all treatments. Leaves on the same phytomer of buds that grew out showed more leaf mass per unit area than those on the same phytomer of buds that remained dormant. CONCLUSIONS: Tillering ceases at specific light conditions within the wheat canopy, independent of population density, and to a lesser extent independent of light intensity. It is suggested that cessation of tillering is induced when the fraction of PAR intercepted by the canopy exceeds a specific threshold (0.40-0.45) and red : far-red ratio drops below 0.35-0.40.     

Simulation of the three-dimensional distribution of the red:far-red ratio within crop canopies

It is widely recognized that the red:far-red ratio (zeta) acts as a signal that triggers plant morphogenesis. New insights into photomorphogenesis have been gained through experiments in controlled environments. Extrapolation of such results to field conditions requires characterization of the zeta signal perceived by plant organs within canopies. This paper presents a modeling approach to characterize this signal. A wheat (Triticum aestivum) architectural model was coupled with a three-dimensional light model estimating the irradiances of virtual sensors. Architectural parameters and zeta values were measured on two contrasting spring wheat canopies under outdoor conditions. Light simulations were compared with measurements, and an analysis of sensitivity to measurement conditions was carried out. The model results agreed well with measurements and previously published data. The sensitivity analysis showed that zeta strongly depends on canopy development as well as on sky conditions, sensor orientation, and sensor field of view. This paper shows that modeling enables investigation of zeta distribution in a canopy over space and time. It also shows that the characterization of light quality strongly depends on measurement conditions, and that any discrepancies in results are likely attributable to different experimental set-ups. The usefulness of this modeling approach for crop photomorphogenesis studies is discussed.     

The regulation of nitrogen accumulation in the grain of wheat plants (Triticum aestivum)

Nitrogen accumulation in the ear of wheat plants ( Triticum aestivum L. cv. Klein Chamaco) during ear growth was studied under 4 experimental conditions. Plants were grown in pots with Perlite or soil, and fertilized with nutrient solutions. In one experiment the plants were grown in a greenhouse and supplied with high (16m M ) or low (1.6 m M ) N in the nutrient solutions until anthesis, and then with or without nitrogen supply until ripening. In a second experiment the plants were grown with high N supply until anthesis, and then for half of the plants light intensity was decreased by 50%, and at the same time. N supply was terminated for half of the plants within each light treatment. A third experiment was similar to the previous one, but was carried out in a growth cabinet under 20% of the maximal irradiance in the greenhouse. In a fourth experiment half the ear was excised at anthesis in half of the plants, and these plants were then supplied with or without nitrogen.
In all experiments there was a linear relation between the rate of N accumulation and the rate of ear growth. A wide range of final individual grain weights and N concentration was observed among the experiments. The same maximum N concentration was observed for all grain sizes, although the N concentration could be different between grains of the same size. The grain N concentration correlated with the rate of N accumulation per unit of ear weight increase during ear growth. It is suggested that in wheat plants there is a dependence of nitrogen transport on carbon transport to the ear, and to the ear, and that the final grain N concentration is determined by the N/C ratio exported from the vegetative tissues.     

Aluminium induces rapid changes in cytosolic pH and free calcium and potassium concentrations in root protoplasts of wheat (Triticum aestivum)

Addition of aluminium chloride (50 μM Al) caused different effects on the transmembrane electrical potential (PD) of root cells in Al-tolerant wheat (Triticum aestivum) cv. Kadett and Al-sensitive cv. WW 20299. As changes in PD of plant cells may depend on transient fluxes of protons, potassium and/or calcium through cell membranes, the effect of Al was investigated on the cytosolic concentrations of these ions in protoplasts isolated from root tips of the same cultivars. The tetra[acetoxymethyl] esters of the fluorescent dyes bis-carboxyethyl-carboxyfluorescein, BCECF, K⁺-binding benzofuran isophthalate, PBFI, and the stilbene chromophore Fura 2-AM were used to determine pH, K⁺ and Ca²⁺, respectively. Changes in fluorescence ratios, directly reflecting changes in [H⁺], [K⁺] and [Ca²⁺] in the cytosol, were determined by photometry fluorescence microscopy. Additions and removals of Al to and from both cultivars caused hyperpolarizations and depolarizations, respectively, but only in the sensitive cv. WW 20299 did the resting PD decrease gradually. Addition of Al to the protoplasts caused rapid changes in cytosolic pH, free [K⁺] and [Ca²⁺]. In both cultivars Al caused a transient oscillating increase in cytosolic [Ca²⁺] for 1 or 2 min and a rapid pH-dependent change in cytosolic [K⁺]. At pH 5 the presence of K⁺ in the medium diminished the Al-induced decrease in cytosolic [K⁺]. Aluminium (50 μM) induced a transient increase in cytosolic [H⁺] (pH decreased) in both cultivars, but the cytosolic pH returned to its initial value only in the Al-tolerant cv. Kadett. In the Alsensitive cv. WW 20299, repeated additions of Al caused a gradual decline in pH. Moreover, in the presence of 1 mM KCl, pH recovered completely in both cultivars. Since only the effect on pH differed in the two cultivars, the more toxic effect of Al on the cv. WW 20299 should be related to the change in pH.     

Salivation into sieve elements in relation to plant chemistry: the case of the aphid Sitobion fragariae and the wheat, Triticum aestivum

Extended sieve element salivation (E1 waveform in the electrical penetration graph) is a characteristic activity during early sieve element punctures, particularly in resistant plants. In order to explore a chemically-mediated mechanism of resistance associated with sieve element salivation, we compared the pattern of feeding behaviour of the aphid, Sitobion fragariae (Walker), on two cultivars of the wheat Triticum aestivum L., with different concentrations of hydroxamic acids (Hx). During 24 h of electronic monitoring, aphids dedicated over 50% of the total time to phloem ingestion from the sieve elements. Total time allocated to E1 in the experiment, time to first E1 within the experiment, time allocated to E1 before a sustained phloem ingestion (E2) and the contribution of sieve element salivation to the phloem phase (E1/[E1+E2]) were significantly higher in the high-Hx cultivar. The increased salivation in plants with higher contents of Hx suggests the existence, at least in this system, of a chemically-mediated sieve element constraint.     

冬小麦(Triticum aestivum)分蘖冗余生态学意义以及减少冗余对水分利用效率的影响

通过盆栽试验,以旱作冬小麦(Triticum aestivum)为材料,分别在拔节和抽穗期对分蘖进行人工干扰,来模拟不可预测的自然干扰,对冬小麦分蘖冗余的生态学意义以及减少这些冗余对水分利用效率影响进行研究.设置3个处理:从拔节期开始剪去所有小的分蘖,仅保留主茎和一个大的分蘖(A);在拔节期剪去主茎和两个大的分蘖,保留所有小的分蘖(B);在孕穗期剪去主茎和有效分蘖,保留无效分蘖(C).没有被干扰的植物作为对照(CK).通过花期测定叶片的叶绿素含量、叶绿素荧光参数、气孔导度和蒸腾速率等生理指标来评价植物的生理与生化活性.结果显示,在拔节期和抽穗期去除主茎和大蘖后,无效分蘖的生理活性被激活,开始执行有效分蘖的功能.到花期时,这些无效分蘖已经在生理活性上满足了补充和替代有效茎的要求.虽然株高和穗的整齐度、穗数和产量显著下降,但并没有防碍小麦的繁衍子代,因此,正是这些由早期"无效分蘖"补充而来的有效茎,避免了小麦绝种的风险.而在拔节期去除无效分蘖后,对小麦产量没有显著影响,但提高了水分利用效率,和对照相比水分利用效率提高了10%.因此,可以认为小麦在分蘖上存在着对水分利用不利的生长冗余,减少这些冗余有望节约用水、提高作物的水分利用效率.     

A comparison of the kinetics of aluminum (Al) uptake and distribution in roots of wheat (Triticum aestivum) using different aluminum sources. A revision of the operational definition of symplastic Al

The lack of information about the movement of aluminum (Al) across the plasma membrane presents a significant barrier to the elucidation of resistance mechanisms which may involve exclusion of Al from the symplast. An understanding of mechanistic aspects of exclusion requires the estimation of symplastic Al levels. Such measurements may be achievable through the use of a kinetic approach. A kinetic protocol was developed to characterize the accumulation and distribution of Al in various cellular compartments in roots of wheat (Triticum aestivum L.). The kinetics of uptake and desorption were similar when Al was supplied as AIK(SO₄)₂ or as AlCl₃. When both salts were supplied at low concentration (50 μM), Al bound to a purified cell wall fraction could be reduced to less than 10–20% of non-exchangeable Al, if roots were washed for 30 min in citric acid following exposure. In contrast, when AlK(SO₄)₂ was supplied at a high concentration (200 μM), a strong linear phase of uptake into cell wall material was observed, which accounted for approximately 48% of non-exchangeable Al in roots. These results suggest that the use of low concentrations of Al in simple salt solutions is required to minimize accumulation of non-exchangeable Al in the apoplasm. A series of multiple-desorption experiments confirmed that citric acid was effective in removing Al from the cell wall compartment of roots exposed to Al for short periods (3 h). However, long exposures (48 h) appeared to create conditions conducive to the accumulation of non-exchangeable Al in the cell wall. In experiments where uptake from solutions containing 50 μM AlCl₃ was followed by desorption in citric acid, non-exchangeable Al in microsomal membrane fractions represented less than 4% of total non-exchangeable Al. Thus, we can exclude the plasma membrane and cell wall as major sites for accumulation of non-exchangeable Al in short exposure studies. Although we cannot provide unequivocal evidence for the localization of Al within the symplast, use of simple salt solutions followed by desorption in citric acid provides the best kinetic technique currently available for the quantitation of Al in the symplasm.     

The impact of photoperiod insensitive Ppd-1a mutations on the photoperiod pathway across the three genomes of hexaploid wheat (Triticum aestivum)

Flowering time is a trait that has been extensively altered during wheat domestication, enabling it to be highly productive in diverse environments and providing a rich source of variation for studying adaptation mechanisms. Hexaploid wheat is ancestrally a long-day plant, but many environments require varieties with photoperiod insensitivity (PI) that can flower in short days. PI results from mutations in the Ppd-1 gene on the A, B or D genomes, with individual mutations conferring different degrees of earliness. The basis of this is poorly understood. Using a common genetic background, the effects of A, B and D genome PI mutations on genes of the circadian clock and photoperiod pathway were studied using genome-specific expression assays. Ppd-1 PI mutations did not affect the clock or immediate clock outputs, but affected TaCO1 and TaFT1, with a reduction in TaCO1 expression as TaFT1 expression increased. Therefore, although Ppd-1 is related to PRR genes of the Arabidopsis circadian clock, Ppd-1 affects flowering by an alternative route, most likely by upregulating TaFT1 with a feedback effect that reduces TaCO1 expression. Individual genes in the circadian clock and photoperiod pathway were predominantly expressed from one genome, and there was no genome specificity in Ppd-1 action. Lines combining PI mutations on two or three genomes had enhanced earliness with higher levels, but not earlier induction, of TaFT1, showing that there is a direct quantitative relationship between Ppd-1 mutations, TaFT1 expression and flowering.     

The significance of the magnesium to manganese ratio in plant tissues for growth and alleviation of manganese toxicity in tomato (Lycopersicon esculentum) and wheat (Triticum aestivum) plants

Results are reported for tomato (Lycopersicon esculentum L. var. Ailsa craig) and wheat (Triticum aestivum L. cv. Mara) which demonstrate that increasing concentrations of Mg in the plant raises plant tolerance to Mn toxicity.Water culture experiments with tomato show that under conditions of high Mn supply (200 µM, Mn), not only does increasing Mg application (0.75 mM to 15 mM) depress Mn uptake, but the higher Mg concentrations in the shoot counteract the onset of Mn toxicity when the concentrations of Mn in the shoot are also high. The ratio of Mg: Mn in the tissues is a better indicator of the appearance of toxicity symptoms than Mn concentration alone. Toxicity symptoms were observed when the Mg:Mn ratio in the shoot tissue was from 1.13 to a value between 3.53 and 6.54. The corresponding Mg: Mn ratio in the older leaves was from 0.82 to between 2.27 and 3.51.For wheat grown in soil, analyses of leaves revealed that growth could be expressed by the following relationship: Y=A+B exp(-kX), where Y=growth, X=Mg:Mn ratio, A, B and k=constants. Growth was significantly reduced when the Mg:Mn ratio fell below 20:1. From a measurement of this ratio it is therefore possible to predict the appearance of Mn toxicity and its influence on growth.     

The effects of elevated atmospheric CO2 on lipid metabolism in leaves from mature wheat (Triticum aestivum cv. Hereward) plants

Winter wheat (Triticum aestivum L. cv. Hereward) plants were grown for 35 d either at 350 μ mol mol^–1 CO₂ or at 650 μ mol mol^–1 CO₂. Lipid synthesis was studied in these plants by incubating the 5th leaf on the main stem with [1–¹⁴C]acetate. Increased CO₂ concentrations did not significantly affect the total incorporation of radiolabel into lipids of whole leaf tissue, but altered the distribution for individual lipid classes. Most noticeable amongst acyl lipids was the reduction in labelling of diacylglycerol and a corresponding increase in the proportion of phosphatidylcholine labelling. In the basal regions, there were similar changes and, in addition, phosphatidylglycerol labelling was particularly increased following growth in an enriched CO₂ atmosphere. The stimulation of labelling of the mitochondrial-specific lipid, diphosphatidylglycerol, prompted an examination of the mitochondrial population in wheat plants. Mitochondria were localized in intact wheat sections by immunolabelling for the mitochondrial-specific chaperonin probe. Growth in elevated CO₂ doubled the number of mitochondria compared to growth in ambient CO₂. Fatty acid labelling was also significantly influenced following growth at elevated CO₂ concentrations. Most noticeable were the changes in 16C:18C ratios for the membrane lipids, phosphatidylcholine, phosphatidylglycerol and monogalactosyldiacylglycerol. These data imply a change in the apportioning of newly synthesized fatty acids between the 'eukaryotic' and 'prokaryotic' pathways of metabolism under elevated CO₂.     

Mode of action of chlorsulfuron in a sensitive wheat (Triticum aestivum) cultivar: primary and secondary effects on nitrogen assimilation

Chlorsulfuron (15 g a.i. ha^-1) inhibited growth of wheat (Triticurn aestivum L. cv. Rongotea) especially on high nitrate (NO₃) supply. Decreased growth at high NO^-₃ was associated with higher concentrations of reduced nitrogen (N) and NO^-₃ in the shoots. Seven days after spraying (DAS), shoot dry weight (dry wt) of sprayed plants was similar with NO^-₃ or branched chain amino acids as main N supply but 28 DAS, shoot dry wt was greater with the amino acid treatment. One DAS, chlorsulfuron caused substantial decreases in extension of the youngest leaf and acetolactate synthase activity and valine content of shoots of plants supplied with NO^-₃ or branched chain amino acids. Total amino acid content of shoots was greater in sprayed plants than in unsprayed plants 1 DAS. Acetolactate synthase activity of sprayed plants supplied low NO^-₃ returned to normal 14–21 DAS. For sprayed plants transferred from low to high NO^-₃ supply 7, 14 or 21 DAS, shoot dry wt 50 DAS increased with increased time of transfer to high NO^-₃ while shoot NO^-₃ content decreased. Shoot NO₃ content of sprayed plants transferred to high NO^-₃ supply 7 or 14 DAS was similar to that in unsprayed plants at applied NO^-₃ concentrations which inhibited growth. It is concluded that inhibition of acetolactate synthase is likely to be the primary mode of action of chlorsulfuron in this wheat cultivar; data are consistent with the proposal that subsequent NO^-₃ accumulation can also inhibit growth.