Inherent trait differences explain wheat cultivar responses to climate factor interactions: New insights for more robust crop modelling

Climate change predictions foresee a combination of rising CO₂, temperature and altered precipitation. Effects of single climatic variables are well defined, but the importance of combined variables and genotypic effects is less known, although pivotal for assessing climate change impacts, for example, with crop growth models. This study provides developmental and physiological data from combined climatic factors for two distinct wheat cultivars (Paragon and Gladius), as a basis to improve predictions for climate change scenarios. The two cultivars were grown in controlled climate chambers in a fully factorial setup of atmospheric CO₂ concentration, growth temperature and watering regime. The cultivars differed considerably in their developmental rate, response pattern and the parameters responsible for most of their variation. The growth of Paragon was linked to climatic effects on photosynthesis and mainly affected by temperature. Paragon was overall more negatively affected by all treatment combinations compared to Gladius. Gladius was mostly affected by watering regime. The cultivars' acclimation strategies to climate factors varied significantly. Thus, considering a single factor is an oversimplification very likely impacting the accuracy of crop growth models. Intraspecific crop variation could help understanding genotype by environment variation. Cultivars with high phenotypic plasticity may have greater resilience against climatic variability.     

A single nucleotide substitution in TaHKT1;5-D controls shoot Na+ accumulation in bread wheat

Improving salinity tolerance in the most widely cultivated cereal, bread wheat (Triticum aestivum L.), is essential to increase grain yields on saline agricultural lands. A Portuguese landrace, Mocho de Espiga Branca accumulates up to sixfold greater leaf and sheath sodium (Na⁺) than two Australian cultivars, Gladius and Scout, under salt stress in hydroponics. Despite high leaf and sheath Na⁺ concentrations, Mocho de Espiga Branca maintained similar salinity tolerance compared to Gladius and Scout. A naturally occurring single nucleotide substitution was identified in the gene encoding a major Na⁺ transporter TaHKT1;5-D in Mocho de Espiga Branca, which resulted in a L190P amino acid residue variation. This variant prevents Mocho de Espiga Branca from retrieving Na⁺ from the root xylem leading to a high shoot Na⁺ concentration. The identification of the tissue-tolerant Mocho de Espiga Branca will accelerate the development of more elite salt-tolerant bread wheat cultivars.     

Changes in plant–soil hydraulic pressure gradients of soybean in response to soil drying

Plant gas‐exchange response to drying soil in many instances tracks a common pattern when expressed as a function of fraction of transpirable soil water (FTSW). There is little decrease in gas exchange until FTSW decreases to a value in the range of about 0.3–0.45, then with further drying gas exchange declines approximately linearly. This unique pattern is hypothesised to reflect mainly changes in the water potential gradient between bulk soil and plant. The primary objective was to directly document the basis of this response by measuring the hydrostatic pressure gradient required in the soil to maintain leaf xylem at zero potential with decreasing FTSW. Pots in which soybean (Glycine max) plants were grown were placed in a pressure chamber and the pressure adjusted to maintain zero water potential in a leaf petiolule. These results showed a small, relatively constant hydrostatic pressure had to be applied to the soil to maintain zero leaf xylem water potential until FTSW decreased to approximately 0.3–0.45. Thereafter, the required hydrostatic pressure gradient increased as FTSW continued to decrease. Hydraulic conductance was calculated to be relatively stable early in the drying cycle, and then decrease as the soil dried to comparatively high FTSW of 0.5–0.7.     

Differential sensitivities of transpiration to evaporative demand and soil water deficit among wheat elite cultivars indicate different strategies for drought tolerance

In order to satisfy increasing wheat demands, scaling up wheat production will require boosting yield in suboptimal, drought-prone areas. Under rain-fed environments, one promising option is the identification of traits allowing for soil water conservation until the next rain episode. This can be achieved either by limiting transpiration rate (TR) of the crop to a maximum level when atmospheric drought (or VPD for vapor pressure deficit) is too high or by decreasing stomata conductance earlier in the soil drying cycle. Although promising, those strategies were never explored in wheat. A first objective of this study was to investigate the extent of the genetic variability of TR sensitivity to both VPD and soil water deficit among a group of eight elite wheat lines, which are cultivated under south Australian conditions. Those consisted of seven differentially drought-adapted lines and one check cultivar. TR responses to VPD were highly variable among genotypes, with six lines displaying a breakpoint in their TR response to VPD that ranged from 2.4 to 3.9 kPa, while two others had their TR increasing linearly as VPD increased. Transpiration response to a progressively decreasing fraction of transpirable soil water (FTSW) was investigated in those lines. A significant genetic variability in the responses among genotypes was observed. They revealed different FTSW thresholds at which transpiration started to decrease at levels ranging from 0.43 to 0.52, and different slopes for the decrease. A second objective was to investigate the existence of phenotypic correlations between the parameters characterizing transpiration sensitivities to both sources of water deficit (i.e., VPD and FTSW). Significant correlations were observed revealing that genotypes with conservative water use in their response to high VPD were also conservative in response to decreasing FTSW and that the drought tolerance of other lines might stem from an apposite strategy, invoking decreased sensitivity of TR to both sources of drought. Those findings provide new options for breeding drought tolerant lines based on this germplasm.     

不同施肥水平对旱地冬小麦水分利用效率的影响

1987—1988年,研究了旱地施肥对冬小麦(Triticum aestivum cv．Shanhe No．6)水分利用效率的影响,初步探讨了“以肥调水”的生理机制。施肥不仅提高了旱地土壤含水量,更重要的是提高了土壤水势和土壤水的有效性,从而增加了有效水分利用。施肥增大旱地冬小麦绿叶面积,延缓叶片衰老,从而降低土壤蒸发,增加蒸腾用水潜势和光合潜势,但净同化率不一定提高。施肥增加旱地冬小麦总的水分利用(ET,即蒸散量)和蒸腾(T)用水,增加地上部生物产量,提高了经济产量和水分利用效率。施肥使冬小麦同时具有耗水和节水以抵御干旱的能力,对植株具有调节作用,使之更好地适应干旱环境。     

Interaction of drought and high temperature on photosynthesis and grain-filling of wheat

Drought and high temperature often occur simultaneously, but their effects on crops are usually investigated individually. Our objective was to compare effects of drought, high temperature, and their interactions on photosynthesis and grain-growth of wheat (Triticum aestivum L.). Plants (cv. Len) were grown uniformly in well-watered soil at 25/20 ± 2 °C day/night until anthesis, when they were subjected to regimes of no drought (soil at field capacity) and drought (plant water potential of –.0 to –2.4 MPa) at 15/10, 25/20, and 35/30 °C in controlled environments until physiological maturity. Drought decreased photosynthesis, stomatal conductance, viable leaf area, shoot and grain mass, and weight and soluble sugar content of kernels but increased plant water-use efficiency. High temperature hastened the decline in photosynthesis and leaf area, decreased shoot and grain mass as well as weight and sugar content of kernels, and reduced water-use efficiency. Interactions between the two stresses were pronounced, and consequences of drought on all physiological parameters were more severe at high temperature than low temperature. The synergistic interactions indicated that productivity of wheat is reduced considerably more by the combined stresses than by either stress alone, and that much of the effect is on photosynthetic processes.     

Photosynthetic Parameters at the Vegetative Stage and during Grain Development of Two Hexaploid Wheat Cultivars Differing in Salt Tolerance

Response of two spring wheat (Triticum aestivum L.) cultivars, salt tolerant SARC-I and salt sensitive Potohar, to different concentrations of NaCl was examined under glasshouse conditions. Eighteen-day-old plants of both the lines grown in sand culture were irrigated with 0 (control), 80, 160 or 240 mM NaCl in full strength Hoagland's nutrient solution. Shoot fresh and dry masses, and leaf area per plant of SARC-I at the vegetative stage, were significantly greater than those of cv. Potohar at higher salt concentrations, however, relative growth rate (RGR) of cv. Potohar was significantly higher than that of SARC-I. SARC-I had higher net photosynthetic rate (P_N), stomatal conductance (g_s) and transpiration rate (E) than cv. Potohar at the vegetative stage, but the cultivars did not differ significantly in water-use efficiency (P_N/E), intrinsic water use efficiency (P_N/g_s), and intercellular/ambient CO₂ concentration ratio. At the grain development stage, SARC-I had significantly higher P_N and g_s in the flag leaf than cv. Potohar under salinity. SARC-I was superior to cv. Potohar with respect to number of grains per spike, number of grains per spikelet, mean grain mass, and grain yield per plant at all NaCl concentrations.     

Physiological mechanisms contributing to the increased water-use efficiency in winter wheat under deficit irrigation

Deficit irrigation in winter wheat has been practiced in the areas with limited irrigation water resources. The objectives of this study were to (i) understand the physiological basis for determinations of grain yield and water-use efficiency in grain yield (WUE) under deficit irrigation; and (ii) investigate the effect of deficit irrigation on dry matter accumulation and remobilization of pre-anthesis carbon reserves during grain filling. A field experiment was conducted in the Southern High Plains of the USA and winter wheat (cv. TAM 202) was grown on Pullman clay loam soil (fine mixed thermic Torretic Paleustoll). Treatments consisted of rain-fed, deficit irrigation from jointing to the middle of grain filling, and full irrigation. The physiological measurements included leaf water potential, net photosynthetic rate (Pn), stomatal conductance (Gs), and leaf area index. The rain-fed treatment had the lowest seasonal evapotranspiration (ET), biomass, grain yield, harvest index (HI) and WUE as a result of moderate to severe water stress from jointing to grain filling. Irrigation application increased seasonal ET, and ET increased as irrigation frequency increased. The seasonal ET increased 20% in one-irrigation treatments between jointing and anthesis, 32-46% in two-irrigation treatments, and 67% in three- and full irrigation treatments. Plant biomass, grain yield, HI and WUE increased as the result of increased ET. The increased yield under irrigation was mainly contributed by the increased number of spikes, and seeds per square meter and per spike. Among the irrigation treatments, grain yield increased significantly but the WUE increased slightly as irrigation frequency increased. The increased WUE under deficit irrigation was contributed by increased HI. Water stress during grain filling reduced Pn and Gs, and accelerated leaf senescence. However, the water stress during grain filling induced remobilization of pre-anthesis carbon reserves to grains, and the remobilization of pre-anthesis carbon reserves significantly contributed to the increased grain yield and HI. The results of this study showed that deficit irrigation between jointing and anthesis significantly increased wheat yield and WUE through increasing both current photosynthesis and the remobilization of pre-anthesis carbon reserves.     

Gas-exchange characteristics,leaf water potential and chlorophyll <Emphasis Type="Italic">a</Emphasis> fluorescence in oil palm (<Emphasis Type="Italic">Elaeis guineensis</Emphasis> Jacq.) seedlings under water stress and recovery

The gas-exchange characteristics, leaf water potential and chlorophyll (Chl) a fluorescence of oil palm (Elaeis guineensis Jacq.) seedlings subjected to water stress and recovery were investigated in a greenhouse experiment. At 24 days after imposition of stress, leaf water potential in water-stressed seedlings was doubled compared to that of control and there was a drastic decline in gas-exchange parameters viz. photosynthesis, transpiration, and stomatal conductance. Water stress did not irreversibly affect gas-exchange parameters and quantum efficiency of photosystem II, as seedlings exhibited total recovery of photosynthetic apparatus by 12^th day of rehydration. These findings indicate that oil palm exhibits physiological plasticity to water stress during the seedling stage.     

The effect of take-all disease on gas-exchange rates and biomass in two winter wheat lines with different drought response

There have been no studies of the effect of take-all on leaf gas-exchange rates, despite the fact that take-all severely restricts plant water and nutrient uptake, which results in significant biomass and grain yield reduction. Here we describe the effect of inoculation with Gaeumannomyces graminis (Sacc.) var. tritici (Ggt) on carbon assimilation rate (A) and biomass production of wheat plants grown under two water regimes. We show that the impact of Ggt inoculation on plant growth and leaf A may be through reduced photosynthetic capacity of the leaves and not water stress per se. The nature of this reduced photosynthetic capacity remains uncertain but may involve nutrient deficiency and different enzymes produced by the fungus. In each of the 3 years the experiment was conducted, Ggt significantly reduced A, i.e. at anthesis by 18% in 2000, 15% in 2001, and 12% in 2002. In agreement with other field studies, Ggt reduced tiller number and production of all plant components, mostly root dry mass and grain mass per plant. Highly significant negative correlations were found between disease rating and A in all years, showing that at disease ratings equal or higher than 3 (on a scale from 1 to 4) A could practically be zero. While A decreased, intercellular CO₂ concentration increased or did not change, and stomatal conductance was relatively high. In addition, A was more reduced under high than under low soil moisture content. These results support the idea that water stress per se did not contribute to the observed reduction of A. The mechanism of photosynthetic capacity reduction due to the Ggt root-rotting fungus is of interest as it may lead to the molecular mechanisms of plant resistance and ultimately to the development of take-all resistant plants.     

Interrelationship between Leaf Gas-Exchange Characteristics,Area Leaf Mass,and Yield in Soybean (Glycine max L. Merr) Genotypes

Variability in leaf gas-exchange traits in thirteen soybean (Glycine max L. Merr) genotypes was assessed in a field experiment conducted at high altitude (1 950 m). Leaf net photosynthetic rate (P ^N) exhibited a high degree of variability at all the growth stages studied. P _N and other gas-exchange parameters exhibited a seasonal pattern that was similar for all the genotypes. P _N rate was highest at seed filling stage. P _N was positively and significantly associated with aboveground dry matter and seed yield. The area leaf mass (ALM) exhibited a strong positive association with leaf P _N, aboveground dry matter, and seed yield. The positive association between ALM, P _N, and seed yield suggests that this simple and easy to measure character can be used in breeding programmes as a surrogate for higher photosynthetic efficiency and eventually higher yield.     

Identification of Differentially Senescing Mutants of Wheat and Impacts on Yield,Biomass and Nitrogen Partitioning

Increasing photosynthetic capacity by extending canopy longevity during grain filling using slow senescing stay-green genotypes is a possible means to improve yield in wheat.Ethyl methanesulfonate (EMS...     

CRISPR/Cas9‐mediated knockout of Ms1 enables the rapid generation of male‐sterile hexaploid wheat lines for use in hybrid seed production

The development and adoption of hybrid seed technology have led to dramatic increases in agricultural productivity. However, it has been a challenge to develop a commercially viable platform for the production of hybrid wheat (Triticum aestivum) seed due to wheat's strong inbreeding habit. Recently, a novel platform for commercial hybrid seed production was described. This hybridization platform utilizes nuclear male sterility to force outcrossing and has been applied to maize and rice. With the recent molecular identification of the wheat male fertility gene Ms1, it is now possible to extend the use of this novel hybridization platform to wheat. In this report, we used the CRISPR/Cas9 system to generate heritable, targeted mutations in Ms1. The introduction of biallelic frameshift mutations into Ms1 resulted in complete male sterility in wheat cultivars Fielder and Gladius, and several of the selected male‐sterile lines were potentially non‐transgenic. Our study demonstrates the utility of the CRISPR/Cas9 system for the rapid generation of male sterility in commercial wheat cultivars. This represents an important step towards capturing heterosis to improve wheat yields, through the production and use of hybrid seed on an industrial scale.     

DIFFERING SELECTION ON PLANT PHYSIOLOGICAL TRAITS IN RESPONSE TO ENVIRONMENTAL WATER AVAILABILITY: A TEST OF ADAPTIVE HYPOTHESES

I used phenotypic selection analysis to test the prediction from functional and comparative studies of plants that smaller leaves and more efficient water use are adaptive in drier environments. I measured selection gradients on leaf size and instantaneous water-use efficiency (a measure of carbon gain per unit water loss) in experimental populations of Cakile edentula var. lacustris placed into wet and dry environments in the field. Linear and nonlinear selection differed significantly between the two environments as predicted. Water-use efficiency was selected to be higher, and leaf area was selected toward a small intermediate optimum, in the dry environment. There was also significant positive correlational selection on water-use efficiency and leaf size, suggesting that the optimum leaf size in the dry environment is greater for plants with higher water-use efficiency. In contrast, neither leaf size nor water-use efficiency were selected in the wet environment, though larger leaves resulted in greater vegetative biomass. Path analysis of the linear selection gradients found that water-use efficiency affected plant fitness primarily because it increased vegetative biomass, as suggested by the hypotheses about the function of physiological traits. These results were not only consistent with the functional hypotheses but also with the observed genetic differentiation in water-use efficiency and leaf size between wet and dry site populations.     

Development of Triticum turgidum subsp. durum - Aegilops longissima amphiploids with high iron and zinc content through unreduced gamete formation in F1 hybrids

Four different interspecific hybrids involving three different accessions of Aegilops longissima Schweinf. & Muschl. with high grain iron and zinc content and three Triticum turgidum L. subsp. durum (Desf.) Husn. cultivars with low micronutrient content were made for durum wheat biofortification and investigated for chromosome pairing, fertility, putative amphiploidy, and micronutrient content. The chromosome pairing in the 21-chromosome F1 hybrids (ABSl) consisted of 0-6 rod bivalents and occasionally 1 trivalent. All the F1 hybrids, however, unexpectedly showed partial but variable fertility. The detailed meiotic investigation indicated the simultaneous occurrence of two types of aberrant meiotic divisions, namely first-division restitution and single-division meiosis, leading to regular dyads and unreduced gamete formation and fertility. The F2 seeds, being putative amphiploids (AABBSlSl), had nearly double the chromosome number (40-42) and regular meiosis and fertility. The F1 hybrids were intermediate between the two parents for different morphological traits. The putative amphiploids with bold seed size had higher grain ash content and ash iron and zinc content than durum wheat cultivars, suggesting that Ae. longissima possesses a better genetic system(s) for uptake and seed sequestration of iron and zinc, which could be transferred to elite durum and bread wheat cultivars and exploited.     

Influence of mulching on the pattern of growth and water use by spring wheat and moisture storage on a fine textured soil

Eight tonnes ha^–1 of stubble were used to mulch spring wheat (Triticum aestivum) on a fine textured soil with the aim of controlling both transpiration and soil evaporation during the wet pre-anthesis phase to increase moisture supply during grain filling in the eastern wheatbelt of Western Australia. Mulching reduced leaf area per plant by reducing the culm number; consequently the green area index was reduced. Reduced culm number was associated with low soil temperature which at 50 mm depth averaged 7°C lower under the mulched crop relative to the control crop in mid-season. The smaller canopies of the mulched crop used 15 mm less water than those of the control before anthesis; this difference in water-use was due equally to reduced transpiration and soil evaporation. However, the mulched crop was unable to increase ET during grain filling, a response associated with the persistence of low soil temperature for most of the growth period. Hence, total ET for the season was significantly lower (18 mm) under the mulched crop than the control crop. At harvest, mulching did not have significant effects on total above-ground dry matter and grain yields, but it increased water use efficiency for grain yield by 18%, grain weight by almost 17% and available moisture in both uncropped and cropped plots by an average of 43 mm.To determine whether there was any residual effects of soil treatment on moisture storage during the summer fallow period, soil moisture was monitored both in cropped plots and uncropped plots, that were either mulched or unmulched during the growing season, from harvest in October 1988 until next planting in June 1989. Available moisture at next planting was correlated with moisture storage at harvest despite the differences in run-off, soil evaporation and fallowing efficiency (increase in moisture storage as a percentage of rainfall) between treatments during fallowing. Therefore, the mulched treatments had more moisture available (30 mm), mostly as a result of less water use during cropping in the previous growing season, than the unmulched treatment.The study shows that mulching may be used to restrain both transpiration and soil evaporation early in the season to increase availability of soil moisture during grain filling. Secondly, mulching during the previous growing season had little effect on soil moisture during the summer fallow period, however, the moisture saved by mulching during cropping was conserved for the following season. These results indicate the importance of evaluating mulching of winter crops in terms of crop yield in the subsequent growing season as well as in the current season in which the soil was treated.Abbreviations D through drainage - DAS days after sowing of the crop on 31 May 1988 - DM dry matter produced in the above-ground portion of the crop (kg ha^–1) - E₀ evaporation from Class A pan (mm) - E_s evaporation from uncropped soil (mm) - E_sc evaporation from soil beneath the wheat canopy (mm) - ET evapotranspiration (mm) - FE fallowing efficiency (gain in soil moisture storage/rainfall) - GAI green area index (area of green vegetation per unit land area) - GWUE water-use efficiency for grain production (grain yield/total ET, kg ha^–1mm^–1) - K extinction coefficient (see equation 1) - RO run-off of moisture from soil surface during/following rainfall (mm) - SM available soil moisture (mm) at harvest (SMh) or at planting (SMp) - WUE water-use efficiency for total above-ground dry matter yield (see GWUE)     

Embryo and endosperm development in wheat (Triticum aestivum L.) kernels subjected to drought stress

The aim of the present work was to reveal the histological alterations triggered in developing wheat kernels by soil drought stress during early seed development resulting in yield losses at harvest. For this purpose, observations were made on the effect of drought stress, applied in a controlled environment from the 5th to the 9th day after pollination, on the kernel morphology, starch content and grain yield of the drought-sensitive Cappelle Desprez and drought-tolerant Plainsman V winter wheat (Triticum aestivum L.) varieties. As a consequence of water withdrawal, there was a decrease in the size of the embryos and the number of A-type starch granules deposited in the endosperm, while the development of aleurone cells and the degradation of the cell layers surrounding the ovule were significantly accelerated in both genotypes. In addition, the number of B-type starch granules per cell was significantly reduced. Drought stress affected the rate of grain filling shortened the grain-filling and ripening period and severely reduced the yield. With respect to the recovery of vegetative tissues, seed set and yield, the drought-tolerant Plainsman V responded significantly better to drought stress than Cappelle Desprez. The reduction in the size of the mature embryos was significantly greater in the sensitive genotype. Compared to Plainsman V, the endosperm cells of Cappelle Desprez accumulated significantly fewer B-type starch granules. In stressed kernels of the tolerant genotype, the accumulation of protein bodies occurred significantly earlier than in the sensitive variety.     

Evidence against the Regulation of Grain Set by Spikelet Abscisic Acid Levels in Water-Stressed Wheat

A possible role of abscisic acid (ABA) in the regulation of grain set in water-stressed wheat (Triticum aestivum L.) was investigated using a split root system to dry half the roots while the remainder were kept watered. Water uptake by the wet roots maintained the leaf water potential at the normal level, whereas the ABA produced in the dry roots was transported to the spike. This caused the spikelet ABA level to increase to the same extent as when the entire root system was stressed to permit a drop in the leaf water potential. In spite of this, the former treatment did not induce a reduction in grain set, whereas the latter did. Thus, contrary to previous reports, water stress-induced changes in spikelet ABA level alone do not appear to regulate grain set.     

Growth and water-use efficiency of 10 Triticum aestivum cultivars at different water availability in relation to allocation of biomass

In environments where the amount of water is limiting growth, water-use efficiency (biomass production per unit water use) is an important trait. We studied the relationships of plant growth and water use efficiency with the pattern of biomass allocation, using 10 wheat cultivars, grown at two soil moisture levels in a growth chamber. Allocation pattern and relative growth rate were not correlated, whereas allocation pattern and water use efficiency were. Variation in transpiration per plant resulted from variation in the rate of transpiration per unit leaf area or root weight, rather than from differences in leaf area or root weight per plant. Transpiration per unit leaf area or root weight was lower when the leaf area or root weight per unit plant weight was larger. Also, the efficiency of water use at the plant and leaf levels was higher for plants with a higher leaf area per unit plant weight, and it was not correlated with the plant's growth rate. Differences in water-use efficiency at the leaf level were related to variation in stomatal conductance, rather than in the rate of photosynthesis. A high photosynthetic water-use efficiency was associated with a low efficiency of nitrogen use for photosynthesis.     

The genetics of nitrogen use in hexaploid wheat: N utilisation, development and yield

A genetic study is presented for traits relating to nitrogen use in wheat. Quantitative trait loci (QTLs) were established for 21 traits relating to growth, yield and leaf nitrogen (N) assimilation during grain fill in hexaploid wheat (Triticum aestivum L.) using a mapping population from the cross Chinese Spring × SQ1. Glutamine synthetase (GS) isozymes and estimated locations of 126 genes were placed on the genetic map. QTLs for flag leaf GS activity, soluble protein, extract colour and fresh weight were found in similar regions implying shared control of leaf metabolism and leaf size. Flag leaf traits were negatively associated with days to anthesis both phenotypically and genetically, demonstrating the complex interactions of metabolism with development. One QTL cluster for GS activity co-localised with a GS2 gene mapped on chromosome 2A, and another with the mapped GSr gene on 4A. QTLs for GS activity were invariably co-localised with those for grain N, with increased activity associated with higher grain N, but with no or negative correlations with grain yield components. Peduncle N was positively correlated, and QTLs co-localised, with grain N and flag leaf N assimilatory traits, suggesting that stem N can be indicative of grain N status in wheat. A major QTL for ear number per plant was identified on chromosome 6B which was negatively co-localised with leaf fresh weight, peduncle N, grain N and grain yield. This locus is involved in processes defining the control of tiller number and consequently assimilate partitioning and deserves further examination. Electronic Supplementary Material The online version of this article () contains supplementary material, which is available to authorized users.