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Stimulation of nodulation in field peas (Pisum sativum) by low concentrations of ammonium in hydroponic culture.

Although the inhibitory effects of high concentrations of mineral N (> 1.0 mM) on nodule development and function have often been studied, the effects of low, static concentrations of NH4+ (< 1.0 mM) on nodulation are unknown. In the present experiments we examine the effects of static concentrations of NH4+ at 0, 0.1 and 0.5 mM in flowing, hydroponic culture on nodule establishment and nitrogenase activity in field peas [Pisum sativum L. cv. Express (Svalöf AB)] for the initial 28 days after planting (DAP). Peas grown in the presence of low concentrations of NH4+ had significantly greater nodule numbers (up to 4-fold) than plants grown without NH4+. Nodule dry weight per plant was significantly higher at 14, 21 and 28 DAP in plants grown in the presence of NH4+, but individual nodule mass was lower than in plants grown without NH4+. The nodulation pattern of the plants supplied with NH4+ was similar to that often reported for supernodulating mutants, however the plants did not express other growth habits associated with supernodulation. Estimates of N2 fixation indicate that the plus-NH4+ peas fixed as much or more N2 than the plants supplied with minus-NH4+ nutrient solution. There were no significant differences in nodule numbers, nodule mass or NH4+ uptake between the plants grown at the two concentrations of NH4+. Nodulation appeared to autoregulate by 14 DAP in the minus-NH4+ treatment. Plant growth and N accumulation in the minus-NH4+ plants lagged behind those of the plus-NH4+ treatments prior to N2 fixation becoming well established in the final week of the experiment. The plus-NH4+ treatments appeared not to elicit autoregulation and plants continued to initiate nodules throughout the experiment.

Culture Media↗

Geometric modeling of inflatable structures for lunar base.

A modular inflatable structure consisting of thin, composite membranes is presented for use in a lunar base. Results from a linear elastic analysis of the structure indicate that it is feasible in the lunar environment. Further analysis requires solving nonlinear equations and accurately specifying the geometries of the structural members. A computerized geometric modeling technique, using bicubic Bezier surfaces to generate the geometries of the inflatable structure, was conducted. Simulated results are used to create three-dimensional wire frames and solid renderings of the individual components of the inflatable structure. The component geometries are connected into modules, which are then assembled based upon the desired architecture of the structure.

Computer Simulation↗

Use of electron microprobe x-ray analysis for determination of low calcium concentrations across leaves deficient in calcium.

An electron microprobe with wavelength-dispersive x-ray spectrometry (WDS) was found to be useful for the determination of Ca concentrations in leaf tissue deficient in Ca. WDS effectively detected Ca concentrations as low as 0.2 mg/g dry wt in the presence of high levels of K and Mg (120 and 50 mg/g dry wt, respectively). Leaf specimens were prepared for analysis by quick-freezing in liquid nitrogen and freeze-drying at -20 degrees C to maintain elemental integrity within the tissue. Because dry material was analyzed, sample preparation was simple and samples could be stored for long periods before analysis. A large beam diameter of 50 gm was used to minimize tissue damage under the beam and analyze mineral concentrations within several cells at one time. Beam penetration was between 50 and 55 microns, approximately one-third of the thickness of the leaf. For analysis of concentrations in interveinal areas, analyses directed into the abaxial epidermis were found most useful. However, because of limited beam penetration, analyses of veinal areas would require use of cross sections [correction of crosssections]. Solid mineral standards were used for instrument standardization. To prevent measurement errors resulting from differences between the matrix of the mineral standards and the analyzed tissue, concentrations in leaves were corrected using gelatin standards prepared and analyzed under the same conditions. WDS was found to be useful for documenting that very low Ca levels occur in specific areas of lettuce leaves exhibiting the Ca deficiency injury termed tipburn.

Calcium↗

Microbial biomass and productivity in seagrass beds.

Different methods for measuring the rates of processes mediated by bacteria in sediments and the rates of bacterial cell production have been compared. In addition, net production of the seagrass Zostera capricorni and bacterial production have been compared and some interrelationships with the nitrogen cycle discussed. Seagrass productivity was estimated by measuring the plastochrone interval using a leaf stapling technique. The average productivity over four seasons was 1.28 +/- 0.28 g C m-2 day-1 (mean +/- standard deviation, n = 4). Bacterial productivity was measured five times throughout a year using the rate of tritiated thymidine incorporated into DNA. Average values were 33 +/- 12 mg C m-2 day-1 for sediment and 23 +/- 4 for water column (n = 5). Spatial variability between samples was greater than seasonal variation for both seagrass productivity and bacterial productivity. On one occasion, bacterial productivity was measured using the rate of 32P incorporated into phospholipid. The values were comparable to those obtained with tritiated thymidine. The rate of sulfate reduction was 10 mmol SO4(-2) m-2 day-1. The rate of methanogenesis was low, being 5.6 mg CH4 produced m-2 day-1. A comparison of C flux measured using rates of sulfate reduction and DNA synthesis indicated that anaerobic processes were predominant in these sediments. An analysis of microbial biomass and community structure, using techniques of phospholipid analysis, showed that bacteria were predominant members of the microbial biomass and that of these, strictly anaerobic bacteria were the main components. Ammonia concentration in interstitial water varied from 23 to 71 micromoles. Estimates of the amount of ammonia required by seagrass showed that the ammonia would turn over about once per day. Rapid recycling of nitrogen by bacteria and bacterial grazers is probably important.

Amines↗

Tomato responses to ammonium and nitrate nutrition under controlled root-zone pH.

Tomato (Lycopersicon esculentum L. Mill. 'Vendor') plants were grown for 21 days in flowing solution culture with N supplied as either 1.0 mM NO3- or 1.0 mM NH4+. Acidity in the solutions was automatically maintained at pH 6.0. Accumulation and distribution of dry matter and total N and net photosynthetic rate were not affected by source of N. Thus, when rhizosphere acidity was controlled at pH 6.0 during uptake, either NO3- or NH4+ can be used efficiently by tomato. Uptake of K+ and Ca2+ were not altered by N source, but uptake of Mg2+ was reduced in NH4(+)-fed plants. This indicates that uptake of Mg2+ was regulated at least partially by ionic balance within the plant.

Biomass↗

Effects of root-zone acidity on utilization of nitrate and ammonium in tobacco plants.

Tobacco (Nicotiana tabacum L., cv. 'Coker 319') plants were grown for 28 days in flowing nutrient culture containing either 1.0 mM NO3- or 1.0 mM NH4+ as the nitrogen source in a complete nutrient solution. Acidities of the solutions were controlled at pH 6.0 or 4.0 for each nitrogen source. Plants were sampled at intervals of 6 to 8 days for determination of dry matter and nitrogen accumulation. Specific rates of NO3- or NH4+ uptake (rate of uptake per unit root mass) were calculated from these data. Net photosynthetic rates per unit leaf area were measured on attached leaves by infrared gas analysis. When NO3- [correction of NO-] was the sole nitrogen source, root growth and nitrogen uptake rate were unaffected by pH of the solution, and photosynthetic activity of leaves and accumulation of dry matter and nitrogen in the whole plant were similar. When NH4+ was the nitrogen source, photosynthetic rate of leaves and accumulation of dry matter and nitrogen in the whole plant were not statistically different from NO3(-) -fed plants when acidity of the solution was controlled at pH 6.0. When acidity for NH4(+) -fed plants was increased to pH 4.0, however, specific rate of NH4+ uptake decreased by about 50% within the first 6 days of treatment. The effect of acidity on root function was associated with a decreased rate of accumulation of nitrogen in shoots that was accompanied by a rapid cessation of leaf development between days 6 and 13. The decline in leaf growth rate of NH4(+) -fed plants at pH 4.0 was followed by reductions in photosynthetic rate per unit leaf area. These responses of NH4(+) -fed plants to increased root-zone acidity are characteristic of the sequence of responses that occur during onset of nitrogen stress.

Biological Transport↗

Root-zone acidity affects relative uptake of nitrate and ammonium from mixed nitrogen sources.

Soybean plants (Glycine max [L.] Merr. cv Ransom) were grown for 21 days on 4 sources of N (1.0 mM NO3-, 0.67 mM NO3- plus 0.33 mM NH4+, 0.33 mM NO3- plus 0.67 mM NH4+, and 1.0 mM NH4+) in hydroponic culture with the acidity of the nutrient solution controlled at pH 6.0, 5.5, 5.0, and 4.5. Dry matter and total N accumulation of the plants was not significantly affected by N-source at any of the pH levels except for decreases in these parameters in plants supplied solely with NH4+ at pH 4.5. Shoot-to-root ratios increased in plants which had an increased proportion [correction of proporiton] of NH4(+)-N in their nutrient solutions at all levels of root-zone pH. Uptake of NO3- and NH4+ was monitored daily by ion chromatography as depletion of these ions from the replenished hydroponic solutions. At all pH levels the proportion of either ion that was absorbed increased as the ratio of that ion increased in the nutrient solution. In plants which were supplied with sources of NO3- plus NH4+, NH4+ was absorbed at a ratio of 2:1 over NO3- at pH 6.0. As the pH of the root-zone declined, however, NH4+ uptake decreased and NO3- uptake increased. Thus, the NH4+ to NO3- uptake ratio declined with decreases in root-zone pH. The data indicate a negative effect of declining root-zone pH on NH4+ uptake and supports a hypothesis that the inhibition of growth of plants dependent on NH4(+)-N at low pH is due to a decline in NH4+ uptake and a consequential limitation of growth by N stress.

Fertilizers↗

Effect of ammonium sulfate, ammonium chloride and root-zone acidity on inorganic ion content of tobacco.

Tobacco plants (Nicotiana tabacum L. cv NC82) were supplied with (NH4)2SO4 or NH4Cl at root-zone pH of 6.0 and 4.5 in hydroponic culture for 28 days. Dry matter accumulation, total N and C content, and leaf area and number were not affected by the NH4+ source or root-zone pH. Plants supplied with NH4Cl accumulated up to 1.2 mM Cl g DW-1, but accumulated 37% less inorganic H2PO4- and 47% less SO4(2-) than plants supplied with (NH4)2SO4. The large Cl- accumulation resulted in NH4Cl- supplied plants having a 31% higher inorganic anion (NO3-, H2, PO4-, SO4(2-), and Cl-) charge. This higher inorganic anion charge in the NH4Cl-supplied plants was balanced by a similar increase in K+ charge. Plants supplied with NH4Cl accumulated greater concentrations of Cl- in leaves (up to 5.1% of DW) than plants supplied with (NH4)2SO4 (less than -% DW). Despite the high Cl- concentration of leaves in NH4Cl supplied plants, these plants showed no symptoms of Cl- toxicity. This demonstrates that toxicity symptoms are not due solely to an interaction between high Cl- concentration in tissue and NH4+ nutrition. The increase in root-zone acidity to pH 4.5 from 6.0 did not induce toxicity symptoms.

Ammonium Chloride↗

Potassium concentration effect on growth, gas exchange and mineral accumulation in potatoes.

This study was conducted to evaluate the responses of potatoes to six K solution concentrations maintained with a flow-through nutrient film system. Potato plants were grown for 42 days in sloping shallow trays containing a 1 cm layer of quartz gravel with a continuous flow of 4 ml min-1 of nutrient solutions having K concentrations of 0.10, 0.55, 1.59, 3.16, 6.44, 9.77 meq L-1. Plant leaf area, total and tuber dry weights were reduced over 25% at 0.10 meq L-1 of K and over 17% at 9.77 meq L-1 of K compared to concentrations of 0.55, 1.59, 3.16 and 6.44 meq L-1 of K. Gas exchange measurements on leaflets in situ after 39 days of growth demonstrated no significant differences among different K treatments in CO2 assimilation rate, stomatal conductance, intercellular CO2 concentration, and transpiration. Further measurements made only on plants grown at 0.10, 1.59, 6.44 meq L-1 of K showed similar responses of CO2 assimilation rate to different intercellular CO2 concentrations. This suggested that the photosynthetic systems were not affected by different K nutrition. The leaves of plants accumulated about 60% less K at 0.10 meq L-1 of K than at higher K concentrations. However, Ca and Mg levels in the leaves were higher at 0.10 meq L-1 of K than at higher K concentrations. This indicates that low K nutrition not only reduced plant growth, but also affected nutrient balance between major cations.

Calcium↗

Growth, carbon dioxide exchange and mineral accumulation in potatoes grown at different magnesium concentrations.

Plants of Norland potatoes (Solanum tuberosum L.) were maintained for 42 days at Mg concentrations of 0.05, 0.125, 0.25, 1, 2, and 4 mM in a nonrecirculating nutrient film system under controlled environment. With the increased Mg supply from 0.05 to 4 mM, Mg concentrations in the leaves of the 42-day old plants increased significantly from 1.1 to 11.2 mg g-1 dry weight. Plant leaf area and plant and tuber dry weights increased with increased Mg concentrations up to 1 mM in solution or 6.7 mg g-1 in leaves, and then decreased with further increases in Mg concentrations. Rates of CO2 assimilation measured on leaflets in situ at ambient and various intercellular CO2 concentrations were consistently lower at 0.05 and 4 mM Mg than at other Mg treatments, which may indicate decreased photosynthetic activity in mesophyll tissues at the lowest and highest Mg concentrations. Dark respiration rates in leaves were highest at 0.05 and 4 mM Mg, lowest at 0.25 and 1 mM Mg, and intermediate at 0.125 and 2 mM Mg. The different Mg treatments also influenced accumulation of other minerals in leaves. Leaf concentrations of Ca and Mn decreased with increased Mg supply except that Ca and Mn were lower at 0.05 mM than at 0.125 mM Mg. Leaf K concentrations were lower at 1, 2 and 4 mM Mg than at other Mg treatments. Foliar concentrations of P, Fe, Zn, and Cu had small but inconsistent variation with different Mg concentrations. Leaf concentrations of N, S, and B were similar at different Mg concentrations. This study demonstrates that various Mg nutrition, along with altered accumulation of other nutrients, could regulate dry matter production in potatoes by affecting not only leaf area but also leaf carbon dioxide assimilation and respiration.

Carbon Dioxide↗

Study of various NH4+/NO3- mixtures for enhancing growth of potatoes.

Two experiments were conducted to determine the effects of various NH4(+)-N/NO3(-)-N percentages on growth and mineral concentrations in potato (Solanum tuberosum L.) plants using a non-recirculating nutrient film system in a controlled environment. The first experiment included six NH4(+)-N/NO3(-)-N percentages at 0/100, 20/80, 40/60, 60/40, 80/20, and 100/0 with the same total N concentration of 4 mM. The second experiment included six NH4(+)-N/NO3(-)-N percentages at 0/100, 4/96, 8/92, 12/88, 16/84, and 20/80 again with the same total N of 4 mM. In each experiment, plants were harvested 35 days after transplanting when tubers had been initiated and started to enlarge. Dry weights of shoots, tubers, and whole plant at the harvest were increased significantly with all mixed nitrogen treatments as compared with single NH4+ or NO3- form. The enhanced growth with mixed nitrogen was greatest at 8% to 20% NH4(+)-N. Also, the concentrations and accumulation of total N in the shoots and roots were greater with mixed nitrogen than with separate NH4+ or NO3- nutrition. With NH4+ present in the solutions, the concentrations of P and Cl in the shoots were increased compared to NO3- alone, whereas the tissue concentrations of Ca and Mg were decreased. It was concluded that nitrogen fertilization provided with combined NH4+ and NO3- forms, even at small proportions of NH4+, can enhance nitrogen uptake and productivity in potato plants.

Dose-Response Relationship, Drug↗

Dry matter and nitrogen accumulation are not affected by superoptimal concentration of ammonium in flowing solution culture with pH control.

While it is known that superoptimal concentrations of the nitrate (NO3-) ion in solution culture do not increase NO3- uptake or dry matter accumulation, the same is not known for the ammonium (NH4+) ion. An experiment was conducted utilizing flowing solution culture with pH control to investigate the influence of superoptimal NH4+ concentrations on dry matter, nitrogen (N), potassium (K), calcium (Ca), and magnesium (Mg) accumulation by nonnodulated soybean plants. Increasing the NH4+ concentration in solution from 1 to 10 mM did not affect dry matter or N accumulation. Accumulations of K, Ca, and Mg were slightly decreased with increased NH4+ concentration. The NH4+ uptake system, which is saturated at less than 1mM NH4+, is able to regulate uptake of NH4+ at concentrations as high as 10 mM.

Biological Transport↗

Changes in ratio of soluble sugars and free amino nitrogen in the apical meristem during floral transition of tobacco.

Under a modification of the nutrient diversion hypothesis, we propose that an inequality in carbohydrate and nitrogen translocation to the apical meristem may be a controlling factor in floral transition. Experiments were conducted in controlled-environment chambers to determine the associations between microscopic characteristics of the transition from vegetative to floral stages of the apical meristem of flue-cured tobacco and to assimilate concentrations in the plant and apical meristem. Low temperature, nitrogen withdrawal, and restriction of nitrogen uptake were used as treatment variables. In all of these stress treatments, flowering occurred at a lesser number of leaves than in control treatments. Low temperature stress accelerated the time of transition to the floral stage as compared with a high temperature control; however, nitrogen stress did not accelerate the time of transition. All stress treatments affected the levels of nitrogen and carbohydrate in whole plants. Most notable was an increase in the percentage of starch and a decrease in the percentage of total soluble carbohydrate induced by the stress treatments. These data indicate that tobacco plants under stress accumulate excess carbohydrate in the form of starch. An apparent inequality in the relative concentrations of carbohydrate and nitrogen in the apical meristem was observed in all treatments at the time of floral transition and is in support of the nutrient diversion hypothesis.

Biological Transport↗

Onset of and recovery from nitrogen stress during reproductive growth of soybean.

Photosynthetic rates and allocation of dry matter, nitrogen, and nonstructural carbohydrates were determined during onset of and recovery from a nitrogen stress for reproductive soybean (Glycine max [L.] Merrill cv Ransom) plants. Until the beginning of seed fill, non-nodulated plants were grown in flowing solution culture with 1.0 mM NO3- in a complete nutrient solution. One set of plants then was transferred to minus-nitrogen solution for 24 d of seed fill; a second set was transferred to a minus-nitrogen solution for 14 d followed by return to the complete solution with 1.0 mM NO3- for the remaining 10 d of seed fill; and a third set was continued on the complete solution. Net CO2 exchange rates of individual leaves, which remained nearly constant during seed fill for nonstressed plants, declined at an accelerated rate during onset of nitrogen stress as the specific content of reduced nitrogen in the leaves was decreased by remobilization of nitrogen to support pod growth. The rate of nitrogen remobilization out of leaves initially was relatively greater than the decrease in photosynthetic rate. While rate of pod growth declined in response to the developing nitrogen stress, photosynthetic assimilation of carbon exceeded reproductive demand and nonstructural carbohydrates accumulated within tissues. Following resupply of exogenous NO3-, specific rate of NO3- uptake by roots was enhanced relative to nonstressed plants. While there was little increase in content of reduced nitrogen in leaves, net remobilization of nitrogen out of leaves ceased, and the decline in photosynthetic rate stabilized at about 51% of that for nonstressed plants. This level of photosynthesis, combined with the availability of elevated pools of carbohydrates accumulated during stress, was sufficient to support the increases in both the specific rates of NO3- uptake and the rate of pod growth during recovery.

Carbohydrate Metabolism↗

Diurnal changes in net uptake rate of nitrate are associated with changes in estimated export of carbohydrates to roots.

The rate of NO3- uptake by soybean (Glycine max [L.] Merrill) roots generally declines during the night in association with progressive depletion of the nonstructural carbohydrate pool in the shoot as well as the concentration of carbohydrates in roots. To determine if NO3- uptake rate changes in response to variations in translocation rate of carbohydrates from shoot to roots per se or to carbohydrate status of the roots, the night period was interrupted with a low light level from incandescent lamps to alter the diurnal pattern of NO3- uptake by roots and export of carbohydrate from shoots of nonnodulated soybean. Depletion of NO3- from replenished, complete nutrient solutions containing 1 mM NO3- was measured by ion chromatography and rates of NO3- uptake were calculated. Changes in export of carbohydrates from shoot to roots during intervals of the night period were calculated as the differences between rates of disappearance in contents of nonstructural carbohydrates and their estimated rates of utilization in shoot respiration and growth. A positive, significant correlation occurred between changes in calculated rates of carbohydrate export from shoots and NO3- uptake rates. Conversely, there was no significant correlation between concentrations of nonstructural carbohydrates in roots and NO3- uptake rates. These results support the hypothesis that carbohydrate flux from shoot to roots has a direct role in regulation of nitrogen uptake by the whole plant.

Biological Transport↗

Internode and petiole elongation of soybean in response to photoperiod and end-of-day light quality.

Elongation of main stem internodes and petioles of soybeans, Glycine max 'Ransom,' was examined in response to various photoperiod/temperature combinations and to end-of-day (EOD) light quality. Photoperiod treatments consisted of 10, 14, and 16 h in combination with day/night temperatures of 18/14, 22/18, 26/22, 30/26, and 34/30 C. The EOD treatments consisted of exposing plants to illumination from either incandescent (high far-red component, FR) or fluorescent (high red component, R) lamps during the final 0.5 h of a 10-h photoperiod. Internode elongation was not significantly promoted by the photoperiod treatments, and, in fact, under the two highest temperature regimes, internode elongation was suppressed under the longer photoperiods. Petiole elongation, however, was enhanced under the longer photoperiods at all temperatures. In the EOD light study, internode and petiole elongation was significantly greater on plants exposed to 0.5 h EOD from incandescent lamps than from fluorescent. Under the incandescent EOD treatment, plants increased dry matter production by 41% and exhibited greater partitioning of assimilates in stem and root portions than under fluorescent EOD.

Environment, Controlled↗

Nitrogen and dry-matter partitioning in soybean plants during onset of and recovery from nitrogen stress.

The study tested the hypothesis that resupplying nitrogen after a period of nitrogen stress leads to restoration of the balance between root and shoot growth and normal functional activity. Nonnodulated soybean plants were grown hydroponically for 14 days with 1.0 mM NO3- in a complete nutrient solution. One set of plants was continued on the complete nutrient solution for 25 days; a second set was given 0.0 mM NO3- for 25 days; and the third set was given 0.0 mM NO3- for 10 days followed by transfer to the complete solution with 1.0 mM NO3- for 15 days. In continuously nitrogen-stressed plants, emergence and expansion of main-stem and branch leaves were severely inhibited as low nitrogen content limited further growth. This was followed by a shift in partitioning of dry matter from the leaves to the roots, resulting in an initial stimulation of root growth and a decreased shoot:root ratio. Reduced nitrogen also was redistributed from the leaves into the stem and roots. When nitrogen stress was relieved, leaf initiation and expansion were renewed. With the restoration of the balance between root and shoot function, the shoot:root ratio and distribution of reduced nitrogen within the plant organs returned to levels similar to those of nonstressed plants.

Biomass↗

Expansion and photosynthetic rate of leaves of soybean plants during onset of and recovery from nitrogen stress.

This study reports on the effects of nitrogen stress and restoration of nitrogen availability after a period of stress on expansion and photosynthetic rate of soybean leaves of differing maturity. We hypothesized that nitrogen resupply would lead to additional accumulation of reduced nitrogen in the leaves and, ultimately, resumption of leaf initiation and expansion and photosynthetic activity. In continuously nitrogen-stressed plants, expansion of middle trifoliolates of main-stem trifoliates and leaf area at full expansion were severely restricted. Leaves showing the greatest effects were initiated after removal of nitrogen. When the reduced nitrogen concentration in mature leaves of continuously stressed plants fell below 9 mg dm-2, the photosynthetic rate per unit leaf decreased rapidly, reaching a minimum of ca. 6-8 mg dm-2 h-1. The older mature leaves tended to abscise at this point, while the youngest leaves remained on the plant and continued to photosynthesize slowly. When nitrogen was resupplied, leaf expansion and final leaf area increased. Leaf initiation was also stimulated as reduced nitrogen levels rose in the leaves. Photosynthetic rates of the oldest and youngest pair of mature leaves returned to values comparable to those of similar-aged leaves of nonstressed soybean plants.

Biomass↗