Nitrogen stress in wheat — Its measurement and relation to leaf nitrogen

Summary The intensity of nitrogen deficiency in the young wheat plant was measured over a range of 1 to 83 per cent of nitrogen stress. Within these limits of tress relative growth rate of tops fell from 11 to 2 per cent per day.Symptoms of nitrogen deficiency appeared when the stress was greater than 40 per cent.Nitrogen stress in the variety Gabo is calibrated in terms of the concentrations of total nitrogen, soluble nitrogen and free ninhydrin nitrogen of the youngest fully expanded leaf.     

Corn (Zea mays L.) growth, leaf pigment concentration, photosynthesis and leaf hyperspectral reflectance properties as affected by nitrogen supply

Plant nitrogen (N)deficiency often limits crop productivity. Early detection of plant N deficiency is important for improving fertilizer N-use efficiency and crop yield. An experiment was conducted in sunlit, controlled environment chambers in the 2001 growing season to determine responses of corn (Zea mays L. cv. 33A14) growth and leaf hyperspectral reflectance properties to varying N supply. Four N treatments were: (1) half-strength Hoagland's nutrient solution applied throughout the experiment (control); (2) 20% of control N starting 15 days after emergence (DAE); (3) 0% N starting 15 DAE; and (4) 0% N starting 23 DAE (0% NL). Plant height, the number of leaves, and leaf lengths were examined for nine plants per treatment every 3–4 days. Leaf hyperspectral reflectance, concentrations of chlorophyll a, chlorophyll b,and carotenoids, leaf and canopy photosynthesis, leaf area, and leaf N concentration were also determined during the experiment. The various N treatments led to a wide range of N concentrations (11 – 48 g kg^–1 DW) in uppermost fully expanded leaves. Nitrogen deficiency suppressed plant growth rate and leaf photosynthesis. At final harvest (42 DAE), plant height, leaf area and shoot biomass were 64–66% of control values for the 20% N treatment, and 46-56% of control values for the 0% N treatment. Nitrogen deficit treatments of 20% N and 0% N (Treatment 3) could be distinguished by changes in leaf spectral reflectance in wavelengths of 552 and 710 nm 7 days after treatment. Leaf reflectance at these two wavebands was negatively correlated with either leaf N (r = –0.72 and –0.75^**) or chlorophyll (r = –0.60 and –0.72^**) concentrations. In addition, higher correlations were found between leaf N concentration and reflectance ratios. The identified N-specific spectral algorithms may be used for image interpretation and diagnosis of corn N status for site-specific N management.     

Photosynthesis and transpiration of the flag leaf in four spring-wheat cultivars

Co₂ exchange and transpiration rates of the flag leaves of four spring wheat (Triticum aestivum L.) cultivars, namely Glenlea, Neepawa, Opal and Kolibri, were compared using infra-red gas-analysis technique. The plants were grown in a controlled environment under an 18-h photoperiod, with day and night temperatures of 20 and 15° C, respectively. The time course of the CO₂-exchange rate (CER) of the flag leaf differed among cultivars. CER began to decrease rapidly some 2 weeks after ear emergence in Glenlea, Neepawa and Kolibri, but only after 4 weeks in Opal. The decline in CER of Glenlea, Neepawa and Opal was continuous throughout the period of grain development whereas in Kolibri CER was maintained at a constant level between the 4th and 6th weeks after ear emergence. The transpiration rates of the flag leaves of the 4 cultivars did not change markedly until 6–7 weeks after ear emergence, indicating that the reduction in CER was not primarily a response to increased stomatal resistance to the diffusion of CO₂. Removing the ear of the main shoot of intact plants failed to depress CER of the subtending flag leaf until 5 weeks after ear removal. Removing the ears of all the tillers of plants in which all but 3 tillers had been removed at ear emergence did not depress CER until 4 weeks after ear emergence, but removal of the ear of the main shoot of plants where all the tillers had been removed at ear emergence reduced the CER of the flag leaf 2 weeks after ear removal. Removal of tillers at ear emergence had a marked effect on the time course of CER and transpiration rates of the flag leaf. Both CER and transpiration rates of a 4-tiller plant were maintained at a higher level throughout ear development as compared to those of a one-tiller plant. The transpiration rate of the flag leaf of Glenlea increased during the later part of the life of the leaf even for one-tiller plants with no ear, indicating that such a stomatal response may be part of the normal course of leaf aging and not a response to a feedback stimulus from the ear.     

低氮诱导玉米幼苗叶片衰老过程中碳氮平衡的动态变化

叶片适时衰老对保证玉米产量有重要意义。本试验以玉米自交系PH6WC和PH4CV为研究对象,通过水培方法,设置低氮(0.04 mmol·L^-1,LN)和正常(4 mmol·L^-1,CK)氮素水平两种处理,在培养2、4、6和8 d后,对其幼苗第2和第3叶片表型、光合特性、叶片中氮素和糖分含量及碳氮比进行分析,旨在探究低氮胁迫下玉米幼苗叶片衰老过程中碳氮平衡的动态变化。结果表明: 与CK相比,LN造成两玉米自交系幼苗第2和第3叶片的面积、生物量、相对叶绿素含量、净光合速率、可溶性糖和淀粉含量均下降,而其氮物质生产能力均先后增加,但第2叶片的变化时间均早于第3叶片;在两叶片的各性状上,均为LN下PH6WC的变化幅度大于PH4CV,且仅幼苗叶片中的碳氮比在LN下显著提高;PH6WC的叶片衰老更快,PH4CV有更强的碳氮平衡能力,其叶片衰老相对滞后。综上,低氮会诱导玉米幼苗叶片衰老,高碳氮比具有促进叶片衰老的调控作用,低氮胁迫下幼苗叶片的碳氮平衡能力在两个玉米基因型间存在较大差异。     

Transpiration of spring barley under nitrogen and phosphorus deficiency during leaf wilting

In an open gas exchange system with a thermocouple psychrometer the transpiration rate of the first leaf in 8-day plants of spring barley was measured in dependence on the water saturation deficit (ΔW _sat). The plants were cultivated in Richter’s nutrient solution, either complete, or deficient in nitrogen or phosphorus. The cuticular transpiration (as measured in the dark) was unaffected by N and P deficiency. The N deficiency reduced the transpiration rate by increasing stomatal resistance since full water saturation of the leaf (67% rate of the control variant) up to stomatal closing at Δ W_sat = 14%. The P deficiency does not affect the transpiration rate at initial phases of wilting, but the stomata close only at a higher Δ W_sat (25%) than those in the control.     

The relation between nitrogen deficiency and second leaf senescence in wheat plants

Life span of the second leaf of wheat(Triticum aestivum L., cv. Grana) plants was studied from day 8 to day 50 of plant age in a variant with nitrogen (+N) and in a variant in which plant senescence was induced by the omission of nitrogen from the nutrient solution (−N). Seed protein was the sole source of nitrogen for these plants. Specific leaf mass (SLM) in the −N variant, and specific leaf area (SLA), the mass of fresh leaf, soluble protein content and total nitrogen content in the +N variant peaked by day 22 of plant age (that is by day 19 of leaf age). Dry matter content, leaf length and leaf area, and SLM in the +N variant peaked by day 29 of plant age (that is by day 26 of leaf age). The ontogeny of the second leaf in the variant with enhanced senescence was shorter by at least 14 days. Plants from this variant showed typical symptoms of N deficiency, that is yellowing of leaves, tip burn, and lack of tillering. However, the growth and biochemical characters studied did not indicate an earlier onset of the senescence of the second leaf of −N plants. Both +N and −N variants reached their peaks (with the exception of an earlier peak by day 12 in case of total nitrogen content in the −N variant) on the same day of leaf age. Thus the first part of the leaf life span from leaf growth initiation to full expansion was of the same length in both the control and N-def icient plants. The stage of the proper senescence of the second leaf of −N plants was very short; the leaf completely died away within 7 days after senescence onset.     

Further evidence for the importance of residual leaf protein complexes in litter decomposition and the supply of nitrogen for plant growth

Summary The nitrogen absorbed by birch seedlings grown in sand culture has been used as a measure of the resistance to decomposition of complexes formed in vitro by interaction between protein and water-extractable leaf constituents ofCalluna vulgaris, Chamaenerion angustifolium andCircaea lutetiana. The resistance to decomposition of these complexes measured in this way isCalluna vulgaris >Chamaenerion angustifolium >Circaea lutetiana. This result seems to be in agreement with the soil-forming and field characteristics of these species and especially the raw humus-forming tendencies ofCalluna vulgaris.The nitrogen released, as measured by the nitrogen absorbed by birch seedlings, fromCalluna raw humus (H-layer material) and the model protein —Calluna leaf extractives complex is very nearly proportional to the widely different amounts of total nitrogen initially added to the root environment thereby suggesting that the organic nitrogen of the model complex and of theCalluna raw humus have similar properties.The amount of nitrogen absorbed by the birch seedlings when unaltered protein was added to the sand culture was less than when protein —Circaea lutetiana leaf extractives complex was added. This is ascribed to too rapid release and loss of nitrogen before it could be absorbed by the birch seedling roots and suggests a role for some leaf protein complexes in the conservation of litter nitrogen. Some possible effects of protein complexes in relation to the digestion of fresh and dry leaf material of various plant species by animals are discussed.Information from archeological observations and from investigations into the tanning of proteins is considered in relation to possible factors influencing the decomposition of protein complexes of litter and it seems probable that a less acid reaction,e.g. such as is brought about by the addition of adequate amounts of calcareous material, will assist the mobilization of the nitrogen of raw-humus-forming litter.The contention that the protein-precipitating substances of leaves are of fundamental importance in soil processes, especially raw humus formation and the supply of nutrients for plant growth, is considered to be supported by experimental approaches from different directions.     

Responses of pest and non-pest Colias butterfly larvae to intraspecific variation in leaf nitrogen and water content

Summary This study examined the effects of intraspecific variation in leaf nitrogen and water content on the growth, consumption, conversion efficiency and nitrogen use of Colias butterfly larvae. Pest and non-pest Colias philodice eriphyle larvae and Colias eurytheme larvae were fed field-collected alfalfa (Medicago sativa) and vetch (Vicia americana) leaves in laboratory experiments. In all treatments, at least one indicator of larval growth performance was positively correlated with leaf nitrogen content, which supports the view that nitrogen is a limiting nutrient for larval growth. The benefits associated with eating leaves with high nitrogen content included higher growth rates, conversion efficiencies, nitrogen accumulation rates and larval nitrogen contents. Over the ranges examined in this study, variation in leaf nitrogen content (2.8–7.0% dry wt) affected larval growth more than variation in leaf water content (66–79% fresh wt). Pest and non-pest C. p. eriphyle responded alike to variation in the leaf nitrogen content of vetch, but there were differences between populations on alfalfa. Pest larvae were more sensitive to variation in leaf water content than non-pest larve. The differences between these populations may be due to specific adaptations resulting from the shift to alfala by pest Colias. It is suggested that herbivores' responses to intraspecific variation in leaf nitrogen content may have important consequences for the evolution of plant defenses and nutrient allocation patterns, and for agricultural pest management.     

Influence of soil applications of nitrogen on nitrate reductase activity,leaf and grain protein content in sorghum

Summary Three sorghum genotypes, L.187, SK5912 and RCFA×L.187 (the latter being a hybrid) were field grown under four nitrogen application rates and replicated four times. The experiments were of complete randomized plot design and conducted in 1976, 1977 and 1978.Nitrate Reductase Activity (NRA) was measured at 5, 7, 9, 11, 13, and 15 weeks and at 18, 19, 20 and 21 weeks after planting in 4th vegetative leaf and flag leaf respectively. Flag leaf, 4th leaf and grain protein contents were also measured.Nitrogen application generally increased NRA in both 4th leaf and flag leaf in the 3 genotypes at all the sampling dates.NRA of the hybrid, RCFA×L.187 was consistently higher than those of SK5912 and L.187 (both straight varieties).NRA of 4th leaf was negatively correlated with leaf protein but flag leaf protein tended to increase with applied nitrogen.Flag leaf NRA was positively and significantly correlated with grain protein indicating an important part played by the flag leaf relative to protein accumulation of the developing grain.     

Effect of N deficiency on leaf growth and cell wall peroxidase activity in contrasting maize genotypes

Two maize genotypes differing in leaf elongation rate (high-LER and low-LER) were used for the investigation of the effects of nitrogen deficiency on leaf growth and development and activity of enzyme cell wall peroxidase in the leaf growth zone. Plants were grown in a growth cabinet in perlite as a substrate and watered with complete N-NO₃ solution (+N) and N-NO₃ deficient solution (–N). Comparison between the investigated genotypes showed that final leaf length in both N treatments was related with LER, but not with the duration of leaf elongation. Faster leaf elongation rate in high-LER compared with low-LER genotype, was associated with longer growth zone, a bigger number of cells in it, and higher cell flux rate, although cell elongation rate was similar in both genotypes. These lines of evidence indirectly indicated that leaves of the faster growing genotype were characterized by higher meristematic activity. Nitrogen deficiency reduced the flux of cells and cell elongation rate, length of cell division zone and the number of cells in whole zone, significantly for both genotypes, although duration of cell elongation was increased and final epidermal cell length was unchanged. These results showed that N deficiency reduced both cell division and cell elongation, which in turn resulted in decreased leaf length and prolonged time for leaf development. Nitrogen deficiency significantly increased both bulk and segmental cell wall peroxidase activity in the growth zone of both investigated genotypes, thus showing an interaction between leaf growth cessation and enzyme activity.     

Differential effects of light and nitrogen on production of hypericins and leaf glands in Hypericum perforatum

The effect of light intensity and root nitrogen supply on the levels of leaf hypericins was examined for St. John’s wort (Hypericum perforatum L.) grown in a sand culture system with artificial lighting. Increasing the light intensity illuminating St. John’s wort plants from 106 to 402 μmol·m^–2·s^–1 resulted in a continuous increase in the level of leaf hypericins. Using a leaf dissection approach, the association of hypericins with the dark glands on the leaves was shown, and it was found that increasing light intensity resulted in a parallel increase in the number of dark glands. In this respect, a linear relationship was observed between leaf gland number and the level of leaf hypericins (R = 0.901). While a decrease in nitrogen supply to St. John’s wort plants also yielded an increase in the level of leaf hypericins, this response occurred in a discontinuous manner over the range of nitrogen levels tested and no significant effect upon the number of dark leaf glands was observed. Overall, these effects of increased light intensity and decreased nitrogen supply on leaf hypericins appear to be independent and additive, and may reflect differences in the sites and processes where these environmental parameters impact production of these phytochemicals.     

Seasonal patterns of growth and nitrogen fixation in field-grown pea

The seasonal patterns of growth and symbiotic N₂ fixation under field conditions were studied by growth analysis and use of¹⁵N-labelled fertilizer in a determinate pea cultivar (Pisum sativum L.) grown for harvest at the dry seed stage. The patterns of fertilizer N-uptake were almost identical in pea and barley (the non-fixing reference crop), but more fertilizer-N was recovered in barley than in pea. The estimated rate of N₂ fixation in pea gradually increased during the pre-flowering and flowering growth stages and reached a maximum of 10 kg N fixed per ha per day nine to ten weeks after seedling emergence. This was the time of early pod-development (flat pod growth stage) and also the time for maximum crop growth rate and maximum green leaf area index. A steep drop in N₂ fixation rate occurred during the following week. This drop was simultaneous with lodging of the crop, pod-filling (round pod growth stage) and the initiation of mobilization of nitrogen from vegetative organs. The application of fertilizer-N inhibited the rate of N₂ fixation only during that period of growth, when the main part of fertilizer-N was taken up and shortly after. Total accumulation of fixed nitrogen was estimated to be 244, 238 and 213 kg N ha⁻¹ in pea supplied with nil, 25 or 50 kg NO ₃ ⁻ −N ha⁻¹, respectively. About one-fourth of total N₂ fixation was carried out during preflowering, one fourth during the two weeks of flowering and the remainder during post-flowering. About 55% of the amount of N present in pods at maturity was estimated to be derived from mobilization of N from vegetative organs. “Starter” N (25 or 50 kg NO ₃ ⁻ −N ha⁻¹) did not significantly influence either dry matter and nitrogen accumulation or the development of leaf area. Neither root length and root biomass determined 8 weeks after seedling emergence nor the yield of seed dry matter and nitrogen at maturity were influenced by fertilizer application.     

In vitro culture of Cuminum cyminum regeneration of flowering shoots from calli of hypocotyl and leaf explants

Calli from hypocotyl explant of Cuminum cyminum L. (Cumin) grew rapidly on Gamborg's B₅ basal medium with the following supplements, (i) 0.5 mg/l — 2,4-D (ii) 4 mg/l — NAA plus 2 mg/l — Kinetin and (iii) 0.2 mg/l — NAA plus 0.2 mg/l — BAP, whereas calli from leaf explant in these media grew slowly. Hypocotyl and leaf calli produced roots when transferred to basal medium only and shoots in basal medium with 0.5 mg/l NAA and 0.1 mg/l BAP. Ninety percent of the shoots produced roots when they were transferred to half strength MS inorganic salts supplemented with 0.5 mg/l each of IBA and NAA.Fifty to sixty percent of rootless as well as rooted shoots produced terminal umbellate flowers on this medium.     

The nature of the organic nitrogen of soils

Summary Examination of the 6N HC1 hydrolysates from 14 different proteins indicated that a considerable proportion of the total protein nitrogen in the hydrolysates, as determined by the micro-Kjeldahl method, was not accounted for by the NH₄-N and the α amino nitrogen found in the hydrolysates. It seems clear that this hydrolysable unidentified nitrogen (HUN) originates mainly from non-amino nitrogen atoms present in arginine, tryptophan, lysine and proline. These nitrogen atoms do not satisfy the conditions necessary for reaction with ninhydrin. The amounts of each amino acid in a particular protein determine the HUN value which will be obtained for 6N HC1 hydrolysates of that protein. There is good agreement between the HUN values for a wide range of proteins when calculated from the amino acid composition of the protein and when determined experimentally. It is concluded that these findings indicate a considerably higher content of amino acid nitrogen in the organic nitrogen of soils and leaf litter than was previously considered to be the case. It is suggested that the findings support the contention that the organic nitrogen of soils contains leaf protein complexes.     

Leaf primordia initiation,leaf emergence and tillering in wheat (Triticum aestivum L.) grown under low-phosphorus conditions

In two simultaneous experiments we examined the effects of phosphorus (P) supply on leaf area development in wheat (Triticum aestivum L.) grown in sand with nutrient solutions. In Experiment 1 we studied leaf emergence, leaf elongation, tiller emergence, shoot growth, and P uptake under four levels of P supply (mM) 0.025 (P1), 0.05 (P2), 0.1 (P3), and 0.5 (P4), and. In Experiment 2 there were two levels of P supply, P1 and P4, and we examined the effects of P on leaf primordia differentiation and leaf emergence. The phyllochron was calculated as the inverse of the rate of leaf emergence calculated from the regression of number of leaf tips (PHY-Ltip), Haun index (PHY-Haun), and as the cumulated thermal time between the emergence of two consecutive leaves (PHYtt). The plastochron was calculated from the inverse of the rate of leaf primordia initiation in the apex. P deficiency delayed the emergence of leaves on the main stem and on the tiller 1. Phosphorus deficiency increased the time from emergence to double ridge and anthesis. The final number of leaves was not affected by P. The effects of P on the value of the phyllochron were attributed to both a reduced rate of leaf primordia initiation, and to a reduced leaf elongation rate. P deficiency delayed or even suppressed the emergence of certain tillers. In this work a phosphorus deficiency that reduced shoot growth by 25% at 44 days after emergence significantly modified the structure of the plants by increasing the value of the phyllochron and delaying tillering. These results suggest that any attempt to simulate leaf area development and growth of wheat plants for P-limited conditions should include the effects of the deficiency on leaf emergence.     

Plant growth, leaf water potential, nitrogenase activity and nodule anatomy in Leucaena leucocephala as affected by water stress and nitrogen availability

 The effects of water stress and nitrogen availability on leaf water potential, nitrogenase activity, and growth was studied in a pot experiment with Leucaena leucocephala seedlings. Water stress was imposed on fertilized and unfertilized plants after inoculation with Rhizobium. Non-inoculated seedlings were used as control plants. Water stress lowered leaf water potential in all seedlings after 14 days of treatment. In inoculated seedlings, fertilized plants were more sensitive to water stress than unfertilized plants, as shown by a higher leaf water potential in plants of the latter treatment. Uninoculated and fertilized seedlings were most affected by water stress. This indicates that Rhizobium might increase stress tolerance in unfertilized seedlings at moderate water stress levels. The combined effects of water stress and applied fertilizers resulted in cessation of nitrogen fixation. Nitrogen fixation came to a complete stop after 22 days of water stress in fertilized seedlings. The different treatments were accompanied by anatomical changes of nodule structure. It is hypothesised that the leaf water potential may be used as an indicator to predict changes in nitrogen fixation in legume tree/shrub species during periods of water stress. Received: 21 October 1996 / Accepted: 12 May 1997     

Nitrogen Deficiency Slows Leaf Development and Delays Flowering in Narrow-leafed Lupin

Effects of nitrogen (N) supply on leaf and flower developmentinLupinus angustifolius L. cv Merrit were examined in a temperature-controlledglasshouse. Low N supply (0.05 or 0.4 mM N) had little effecton leaf initiation but slowed leaf emergence on the main stemcompared with plants receiving high N supply (6.0 or 6.4 mMN), or with symbiotic N₂-fixation. Plants experiencing transientN deficiency had slower leaf emergence than plants with a continuoussupply of 6.4 mM N. Nitrogen supply did not affect the timeof floral initiation, which occurred within 4 weeks of sowing,by which time nine to ten leaves had emerged. However, the floweringof low-N plants was delayed by 68 to 220 °C d (i.e. 4–14d) even though they had fewer leaves. The effect of N deficiencyon flowering time was largely a result of slower leaf emergence. Lupinus angustifolius L.; nitrogen; leaf; flower initiation; thermal time; plastochron; phyllochron     

Nitrogen deficiency inhibits leaf blade growth in Lolium perenne by increasing cell cycle duration and decreasing mitotic and post-mitotic growth rates

Nitrogen deficiency severely inhibits leaf growth. This response was analysed at the cellular level by growing Lolium perenne L. under 7.5 mM (high) or 1 mM (low) nitrate supply, and performing a kinematic analysis to assess the effect of nitrogen status on cell proliferation and cell growth in the leaf blade epidermis. Low nitrogen supply reduced leaf elongation rate (LER) by 43% through a similar decrease in the cell production rate and final cell length. The former was entirely because of a decreased average cell division rate (0.023 versus 0.032 h(-1)) and thus longer cell cycle duration (30 versus 22 h). Nitrogen status did not affect the number of division cycles of the initial cell's progeny (5.7), and accordingly the meristematic cell number (53). Meristematic cell length was unaffected by nitrogen deficiency, implying that the division and mitotic growth rates were equally impaired. The shorter mature cell length arose from a considerably reduced post-mitotic growth rate (0.033 versus 0.049 h(-1)). But, nitrogen stress did not affect the position where elongation stopped, and increased cell elongation duration. In conclusion, nitrogen deficiency limited leaf growth by increasing the cell cycle duration and decreasing mitotic and post-mitotic elongation rates, delaying cell maturation.     

Effect of specific leaf nitrogen content on photosynthesis of sugarcane

In the laboratory, rate of photosynthesis (P) of the youngest fully emerged leaf lamina on a shoot, whether taken from a 4 or 8 month old sugarcane plant, increased approximately linearly with specific leaf nitrogen content (SLN) from < 1.0 to c. 1.7 g m^-2; the rate of increase was greater the higher the incident photosynthetically active radiation (PAR) flux density, over the range 80 to 1600 μmol m^-2s^-1. In a field crop receiving standard fertiliser, including 150 kg N ha^-1, SLN of the green leaf canopy decreased more or less linearly with depth, from > 1.7 g m^-2at the top to c. 1.0 g m^-2at the bottom at 15 wk of age, and progressively with time to 1.2 to 0.85 g m^-2from top to bottom of the canopy by week 45. Where an extra 300 N ha^-1were applied in top-dressings, the decrease in SLN with time was somewhat smaller, to 1.3 to 0.9 g m^-2from top to bottom of the canopy by week 45. Calculations with a model, using the data from laboratory and field, suggest a decrease of about 25% in canopy gross P and about 30% in canopy net P from wk 15 to wk 45 in the crop receiving standard fertiliser only; the calculated decreases where the crop received additional nitrogen were about 17% and 20% respectively. There would seem to be a need for a change in type of cultivar to make profitable use of augmented nitrogen fertiliser.     

Uptake and distribution of nitrogen in wheat plants supplied with different amounts of nitrogen after stem elongation

The uptake of nitrogen and its distribution between shoots and between organs within shoots in wheat (Triticum aestivum) was studied from the start of stem elongation to 28 days after anthesis in a glasshouse experiment with eight nitrogen levels between 0·1 and 12·8 mequiv./litre. There was no net uptake of nitrogen in plants supplied with 0·8 mequiv./litre or less; with more nitrogen the absorption increased linearly. Twenty to 44% of the total plant nitrogen was absorbed after anthesis, this fraction increasing with nutrient supply. The nitrogen allotted to the main shoot decreased until the onset of anthesis and increased thereafter at the expense of the tillers, except with 12·8 mequiv./litre, where nitrogen percentage in the main shoot decreased also after anthesis. Raising nitrogen supply increased the proportion of plant nitrogen recovered in the tillers. Nitrogen accumulated in the ear after emergence and by the 28th day after anthesis it contained between 52% and 73% of the total plant nitrogen. The ear of the main shoot had a higher proportion of shoot nitrogen than that of the tillers. The fraction of ear nitrogen supplied by retranslocation decreased from almost 100% with 0·8 mequiv. N/litre or less to nil with 12·8 mequiv./litre. Increasing nitrogen application decreased the fraction of total nitrogen allocated to the ear.