Nitrogen Utilization in N-limited Barley During Vegetative and Generative Growth: IV. TRANSLOCATION AND REMOBILIZATION OF NITROGEN

Barley (Hordeum vulgare L., cvs Golf and Laevigatum) was grownunder nitrogen limitation in solution culture until near maturity.Three different nitrogen addition regimes were used: in the‘HN’ culture, the relative rate of nitrate-N additionwas 0·08 d^–1 until day 48 and then stepwise decreasedto, finally, 0·005 d^–1 during late grain-filling;the ‘LN’ culture received 45% of the nitrogen addedin HN; the ‘CN’ culture was maintained at RA 0·0375d^–1 throughout growth. At four different growth stages(vegetative,anthesis, and twice during grain-filling), ¹⁵N-nitrate was fedto the plants. In some cases (‘split root cultures’),label was fed only to one-half of the root system. These wereharvested directly after labelling, whereas ‘standardcultured’ plants were harvested at termination of theexperiment (day 148). Absorption of added nitrate was nearlycomplete in the HN and LN cultures, and translocation of nitrogenwithin the plants could thus be studied independently of differencesin nitrate absorption. Cycling of nitrogen absorbed by vegetativeplants accounted for up to 50% of the nitrogen recovered inthe roots. The sink strength of the roots for cycling nitrogen,however, declined during post-anthesis growth, and net lossof nitrogen from both roots and vegetative shoot tissue occurredconcomitantly with incorporation of labelled ¹⁵N-nitrogen. Thenitrogen of the vegetative shoot tissue was substantially lesslabelled than the nitrogen entering the ears, indicating thattranslocation of recently absorbed nitrogen to ears occurs withminor prior exchange with the bulk nitrogen of shoots. In caseswhere the sink strength of the ears was weak, as in LN-culturedLaevigatum (due to high frequency of sterile flowers) and inCN-cultured Golf, nitrogen translocated from roots appearedto be incorporated into the vegetative shoot tissue. There werealso indications that a fraction of the remobilized nitrogenwas actually lost from the plants in these cases. It is concludedthat the root remains efficient in translocation of nitrogento the aerial parts throughout ontogeny and that nitrogen takenup during grain–filling is preferentially directly translocatedto the developing grains. The further translocation of nitrogenreceived by vegetative shoot parts to ears appears mainly relatedto the potential of the ear to accumulate nitrogen. Nitrogenabsorbed/remobilized in excess of the sink strength of the earsis either invested in continued shoot growth, or is irreversiblylost from the plants. Key words: Barley, ¹⁵N-labelling, post-anthesis, remobilization, translocation     

Nitrogen Utilization in N-limited Barley During Vegetative and Generative Growth: II. METHOD FOR MONITORING GENERATIVE GROWTH AND DEVELOPMENT IN SOLUTION CULTURE

Growth of barley (Hordeum vulgare L., cvs Golf, Mette, and Laevigatum)under N-limitation was investigated in solution culture, withspecial emphasis on the generative growth stage. Three differentregimes for limiting nitrate-N availability while keeping otherelements in surplus were employed. In the ‘high, decreasingnitrogen’ (HN) treatment, the relative nitrogen additionrate (RA) was maintained at 0.08 d^–1 until the ears startedto develop and was then decreased stepwise to, finally, RA 0·005d^–1. In the ‘low, decreasing nitrogen’ (LN)treatment, plants received 45% of the nitrogen added in HN.In the ‘constant nitrogen’ (CN) treatment, RA washeld constant at 0·0375 d^–1. Cumulative nitrogenadditions at termination of the experiment (day 147 after sowing)were 192, 179 and 87 mg plant^–1 for HN, CN, and LN cultures,respectively. Nitrogen availability limited nitrogen acquisitionin all treatments except in the CN culture at plant ages above110 d. Stepwise decrease of RA largely synchronized transitionsin developmental stages in the different cultivars and nitrogenregimes (HN and LN), and eventually yielded plants that wereclose to completing their life cycle. Normal maturation wasnot obtained in the CN treatment. The HN and LN treatments wereused for formal analysis of post-anthesis growth. A sigmoidfunction was fitted to growth data and from this, organ weightsand nitrogen concentrations at maturity (defined as cessationof growth) were derived. The two modern cultivars, Golf andMette, clearly outperformed the more primitive cv. Laevigatumin terms of allocation of nitrogen to ears, particularly inthe LN culture. The stepwise decrease of RA appears suitablefor studies of post-anthesis growth and nitrogen relations inbarley, with regard to both genotypic variation and variationcaused by differences in cumulative amounts of nitrogen added. Key words: Barley, development, growth, nitrogen concentration, post-anthesis     

Nutrient Supply and the Growth of the Seminal Root System in Barley: II. LOCALIZED, COMPENSATORY INCREASES IN LATERAL ROOT GROWTH AND RATES OP NITRATE UPTAKE WHEN NITRATE SUPPLY IS RESTRICTED TO ONLY PART OF THE ROOT SYSTEM

The development of the seminal root system, its ability to absorbnitrate, and effects on shoot growth were studied in barleyplants in nutrient solution. The roots received either a uniformsupply of 1.0 mM nitrate (controls), or a supply of the samesolution restricted to a 4-cm length of only one of the mainseminal roots (axes) on each plant, the remainder of the rootsystem receiving a solution containing a low concentration (0.01mM). Marked increases took place in both the growth of lateralroots and the absorption and transport of ¹⁵N-labelled nitrate(per unit root weight) from the zone locally supplied with 1.0mM nitrate. These effects appear largely to compensate for thedeficient supply of nitrate to the remainder of the root system,since after 14 d the relative growth rate (g g^–1d^–1)of the total plant equals that of the controls. Rates of ¹⁵N-nitrateuptake (per unit root weight) remain relatively uniform throughoutthe 29-d experiment, during which root axes develop from theirinitial unbranched form to a complex system of laterals. Theresults are discussed in relation to possible mechanisms bywhich coordination is maintained between root growth, ion uptake,and shoot growth.