Uptake of ammonium and nitrate by carob (Ceratonia siliqua) as affected by root temperature and inhibitors

Seedlings of carob ( Ceratonia siliqua L. cv. Mulata) were grown in nutrient solution culture for 5 weeks, with or without nitrogen at different root temperatures (10, 16, 22, 30, 35 or 40deg;C) and with the air temperature kept between 20 and 24°C. The nitrogen was given as either ammonium or nitrate. At all root temperatures studied, nitrogen-depleted plants developed higher net uptake rates for nitrogen than plants grown in the presence of nitrogen. Temperature affected the kinetic parameters of nitrate uptake more than those of ammonium uptake. With increasing root temperature, the K_m of ammonium uptake decreased, but to a lesser extent than the K_m for nitrate. The increase in V_max of ammonium uptake with temperature was also less noticeable than that for nitrate uptake. Ammonium and nitrate uptakes were inhibited in a similar way by respiratory or protein synthesis inhibitors. It may be noted that ammonium uptake in the presence of inhibitors at 40°C was higher than uptake at 10°C without inhibitors. Some similarities between the transport mechanisms for nitrate and ammonium are underlined in the present work. Components of both transport systems displayed saturation kinetics and depended on protein synthesis and energy. The following components of nitrate uptake were distinguished: (a) a passive net influx into the apparent free space; (b) a constitutive active uptake and (c) active uptake dependent on protein synthesis. We may similarly define three ammonium uptake systems: (a) a passive influx into the apparent free space; (b) passive diffusion uptake at high temperature and (c) active uptake dependent on protein synthesis. The possible role of the ratio between mechanism (c) and mechanism (b) as determinant of ammonium sensitivity is discussed.     

Effect of ammonium or nitrate nutrition on net photosynthesis,growth, and activity of the enzymes nitrate reductase and glutamine synthetase in blueberry,raspberry and strawberry

Blueberry, raspberry and strawberry may have evolved strategies for survival due to the different soil conditions available in their natural environment. Since this might be reflected in their response to rhizosphere pH and N form supplied, investigations were carried out in order to compare effects of nitrate and ammonium nutrition (the latter at two different pH regimes) on growth, CO2 gas exchange, and on the activity of key enzymes of the nitrogen metabolism of these plant species. Highbush blueberry (Vaccinium corymbosum L. cv. 13–16–A), raspberry (Rubus idaeus L. cv. Zeva II) and strawberry (Fragaria × ananassa Duch. cv. Senga Sengana) were grown in 10 L black polyethylene pots in quartz sand with and without 1% CaCO3 (w: v), respectively. Nutrient solutions supplied contained nitrate (6 mM) or ammonium (6 mM) as the sole nitrogen source. Compared with strawberries fed with nitrate nitrogen, supply of ammonium nitrogen caused a decrease in net photosynthesis and dry matter production when plants were grown in quartz sand without added CaCO3. In contrast, net photosynthesis and dry matter production increased in blueberries fed with ammonium nitrogen, while dry matter production of raspberries was not affected by the N form supplied. In quartz sand with CaCO3, ammonium nutrition caused less deleterious effects on strawberries, and net photosynthesis in raspberries increased as compared to plants grown in quartz sand without CaCO3 addition. Activity of nitrate reductase (NR) was low in blueberries and could only be detected in the roots of plants supplied with nitrate nitrogen. In contrast, NR activity was high in leaves, but low in roots of raspberry and strawberry plants. Ammonium nutrition caused a decrease in NR level in leaves. Activity of glutamine synthetase (GS) was high in leaves but lower in roots of blueberry, raspberry and strawberry plants. The GS level was not significantly affected by the nitrogen source supplied. The effects of nitrate or ammonium nitrogen on net photosynthesis, growth, and activity of enzymes in blueberry, raspberry and strawberry cultivars appear to reflect their different adaptability to soil pH and N form due to the conditions of their natural environment. This revised version was published online in July 2006 with corrections to the Cover Date.     

Importance of seed Zn content for wheat growth on Zn-deficient soil

Eggplants (Solanum melongena L. cv. Bonica) were grown in a glasshouse during summer under natural light with one unbranched shoot or one shoot with 3 to 4 branches and with or without fruit in quartz sand buffered and not buffered with 0.5% CaCO₃ (w : v), respectively. Nutrient solutions supplied contained nitrate or ammonium as the sole nitrogen source. Compared with nutrient solutions containing nitrate (10 mM), solutions containing ammonium (10 mM) caused a decrease in net photosynthesis of eggplants during early stages of vegetative growth when grown in quartz sand not buffered with CaCO₃. The decrease was not observed before leaves showed interveinal chlorosis. In contrast, net photosynthesis after bloom at first increased more rapidly in eggplants supplied with ammonium than with nitrate nitrogen. However, even in this case, net photosynthesis decreased four weeks later when ammonium nutrition was continued. The decrease was accompanied by epinasty and interveinal chlorosis on the lower leaves and later by severe wilting, leaf drop, stem lesions, and hampered growth of stems, roots, and fruits. These symptoms appeared later on plants not bearing fruits than on plants bearing fruits. If nutrient solutions containing increasing concentrations of ammonium (0.5–30 mM) were supplied after the time of first fruit ripening, shoot growth and set of later flowers and fruits were promoted. In contrast, vegetative growth and reproduction was only slightly affected by increasing the concentration of nitrate in the nutrient solutions. In quartz sand buffered with CaCO₃ ammonium nutrition caused deleterious effects only under low light conditions (shade) and on young plants during rapid fruit growth. If eggplants were supplied with ammonium nitrogen before bloom, vegetative growth was promoted, and set of flowers and fruit occurred earlier than on plants supplied with nitrate. Furthermore, the number of flowers and fruit yield increased. These effects of ammonium nutrition were more pronounced when plants were grown with branched shoots than with unbranched shoots. The results indicate that vegetative and reproductive growth of eggplants may be manipulated without causing injury to the plants by supplying ammonium nitrogen as long as the age of the plants, carbohydrate reserves of the roots, quantity of ammonium nitrogen supplied, and pH of the growth medium are favourable. T W Rufty Section editor     

不同氮素形态及水分胁迫对水稻苗期水分吸收、光合作用及生长的影响

采用室内营养液培养, 聚乙二醇(PEG6000)模拟水分胁迫处理、HgCl2抑制水通道蛋白活性的方法, 在3种供氮形态下(NH4+-N/ NO 3--N为100/0、50/50和0/100), 研究了水稻苗期水分吸收、光合及生长的状况。结果表明, 在非水分胁迫下, 水稻单位干重吸水量以单一供NO3--N处理最高, 加HgCl2抑制水通道蛋白活性后, 单一供NO3--N、NH4+-N和NH4+-N/ NO3--N为50/50处理的水稻水分吸收分别下降了9.6%、20.7%和16.0%; 但在水分胁迫下, 单一供NO3--N的处理水分吸收量显著降低, 低于其它2个处理, 加HgCl2抑制水通道蛋白活性后, 水分吸收量分别降低了1.0%、18.8%和23.5%。在2种水分条件(水分胁迫与非水分胁迫)下, 净光合速率、气孔导度、蒸腾速率和细胞间隙CO2浓度等指标均以单一供NH4+-N处理最大,NH4+-N/ NO3--N为50/50处理次之, 单一供NO3--N处理最小。HgCl2处理结果表明, 不同形态氮素营养能够影响水稻幼苗根系水通道蛋白活性。在2种水分条件下, NH4+-N/ NO3--N为50/50处理的生物量(干重)均最大。本研究为水稻苗期合理施肥以壮苗提供了理论依据。     

不同氮素形态及水分胁迫对水稻苗期水分吸收、光合作用及生长的影响

采用室内营养液培养,聚乙二醇（PEG6000）模拟水分胁迫处理、HgCl2抑制水通道蛋白活性的方法,在3种供氮形态下（NH4^＋-N／NO36-N为100／0、50／50和0／100）,研究了水稻苗期水分吸收、光合及生长的状况。结果表明,在非水分胁迫下,水稻单位干重吸水量以单一供NO3^--N处理最高,加HgCl2抑制水通道蛋白活性后,单一供NO3^--N、NH4^＋-N和NH4^＋-N／NO3^--N为50,50处理的水稻水分吸收分别下降了9．6％、20．7％和16．0％;但在水分胁迫下,单一供N03^--N的处理水分吸收量显著降低,低于其它2个处理,加HgCl2抑制水通道蛋白活性后,水分吸收量分别降低了1．0％、18．8％和23．5％。在2种水分条件（水分胁迫与非水分胁迫）下,净光合速率、气孔导度、蒸腾速率和细胞间隙CO2浓度等指标均以单一供NH4^＋-N处理最大,NH4^＋-N／NO3^--N为50,50处理次之,单一供NO3^--N处理最小。HgCl2处理结果表明,不同形态氮素营养能够影响水稻幼苗根系水通道蛋白活性。在2种水分条件下,NH4^＋-N／N03^--N为50,50处理的生物量（干重）均最大。本研究为水稻苗期合理施肥以壮苗提供了理论依据。     

Effects of Exposure to Ammonium and Transplant Shock upon the Induction of Nitrate Absorption

In barley (Hordeum vulgare L. cv Steptoe) seedlings, the time course for induction of root nitrate absorption varied significantly with pretreatment. Net nitrate uptake of nitrogen-deprived plants more than doubled during the 12 hours after first exposure to nitrate. For these plants, gentle physical disturbance of the roots inhibited net nitrate absorption for more than 6 hours and potassium absorption for 2 hours. Pretreatment with ammonium appeared sufficient to induce nitrate absorption; plants either grown for 2 weeks on or exposed for only 10 hours to a medium containing ammonium as a sole nitrogen source showed high rates of net nitrate uptake when first shifted to a medium containing nitrate. Gentle physical manipulation of these plants inhibited nitrate absorption for 2 hours and potassium absorption for more than 12 hours. These results indicate (a) that experimental protocols should avoid physical manipulation of the roots when-ever possible and (b) that ammonium or a product of ammonium assimilation can induce nitrate absorption.     

Functional aspects of root architecture and mycorrhizal inoculation with respect to nutrient uptake capacity

The aim of this research was to investigate the effect of arbuscular mycorrhizal (AM) colonisation on root morphology and nitrogen uptake capacity of carob ( Ceratonia siliqua L.) under high and low nutrient conditions. The experimental design was a factorial arrangement of presence/absence of mycorrhizal fungus inoculation ( Glomus intraradices) and high/low nutrient status. Percent AM colonisation, nitrate and ammonium uptake capacity, and nitrogen and phosphorus contents were determined in 3-month-old seedlings. Grayscale and colour images were used to study root morphology and topology, and to assess the relation between root pigmentation and physiological activities. AM colonisation lead to a higher allocation of biomass to white and yellow parts of the root. Inorganic nitrogen uptake capacity per unit root length and nitrogen content were greatest in AM colonised plants grown under low nutrient conditions. A better match was found between plant nitrogen content and biomass accumulation, than between plant phosphorus content and biomass accumulation. It is suggested that the increase in nutrient uptake capacity of AM colonised roots is dependent both on changes in root morphology and physiological uptake potential. This study contributes to an understanding of the role of AM fungi and root morphology in plant nutrient uptake and shows that AM colonisation improves the nitrogen nutrition of plants, mainly when growing at low levels of nutrients.     

氮素形态对小白菜生长和碳氮积累的影响

水培条件下,研究不同氮素形态(硝态氮、铵态氮、甘氨酸、谷氨酰胺、丙氨酸、牛血清蛋白,以及甘氨酸与硝态氮、牛血清蛋白与硝态氮的混合氮源)对小白菜生长和碳氮积累的影响.结果表明:不同氮素形态对小白菜质量、碳氮积累量、可溶性蛋白质含量、可溶性糖含量和游离氨基酸含量的影响不同;硝态氮处理下小白菜地上部分和根的干质量与鲜质量均最大;甘氨酸对小白菜根系的生长及碳氮积累具有明显的促进作用;在3种氨基酸中,谷氨酰胺更有利于小白菜地上部分的生长和氮积累.聚类分析表明,9种氮素形态处理按营养效应大小分为:硝态氮、谷氨酰胺＞甘氨酸与硝态氮混合氮源、牛血清蛋白与硝态氮混合氮源、甘氨酸、铵态氮＞丙氨酸、牛血清蛋白、对照.有机氮源可以作为小白菜生长的氮源,不同的氮素形态对植物产生的生理效应不同.     

The effect of nitrogen source on photosynthesis of carob at high CO2 concentrations

Carob seedlings ( Ceratonia siliqua L. cv. Mulata), fed with nitrate or ammonium, were grown in growth chambers containing two levels of CO₂ (360 or 800 μl l⁻¹), three root temperatures (15, 20 or 25°C), and the same shoot temperature (20/24°C, night/day temperature). The response of the plants to CO₂ enrichment was affected by environmental factors such as the type of inorganic nitrogen in the medium and root temperature. Increasing root temperature enhanced photosynthesis rate more in the presence of nitrate than in the presence of ammonium. Differences in photosynthetic products were also observed between nitrate- and ammonium-fed carob seedlings. Nitrate-grown plants showed an enhanced content of sucrose, while ammonium led to enhanced storage of starch. Increase in root temperature caused an increase in dry mass of the plants of similar proportions in both nitrogen sources. The enhancement of the rates of photosynthesis by CO₂ enrichment was proportionally much larger than the resulting increases in dry mass production when nitrate was the nitrogen source. Ammonium was the preferred nitrogen source for carob at both ambient and high CO₂ concentrations. The level of photosynthesis of a plant is limited not only by atmospheric CO₂ concentration but also by the nutritional and environmental conditions of the root.     

Constrained preferences in nitrogen uptake across plant species and environments

Knowledge of determining factors for nitrogen uptake preferences and how they are modified in changing environments are critical to understand ecosystem nitrogen cycling and to predict plant responses to future environmental changes. Two ¹⁵N tracer experiments utilizing a unique differential labelled nitrogen source were employed in both African savannas and greenhouse settings. The results demonstrated that nitrogen uptake preferences were constrained by the climatic conditions. As mainly indicated by root δ¹⁵N signatures at 1:1 ammonium/nitrate ratio, in the drier environments, plants preferred nitrate and in the wetter environments they preferred ammonium. Nitrogen uptake preferences were different across different ecosystems (e.g. from drier to wetter environments) even for the same species. More significantly, our experiments showed that the plant progeny continued to exhibit the same nitrogen preference as the parent plants in the field, even when removed from their native environment and the nitrogen source was changed dramatically. The climatic constraint of nitrogen uptake preference is likely influenced by ammonium/nitrate ratios in the native habitats of the plants. The constancy in nitrogen preference has important implications in predicting the success of plant communities in their response to climate change, to seed bank use and to reforestation efforts.     

局部根区水分胁迫下氮形态与供给部位对玉米幼苗生长的影响

通过向玉米幼苗分根装置一侧根室的营养液中加入聚乙二醇(PEG 6000)来模拟植物水分胁迫,并设3种供氮形态(硝态氮、铵态氮、两者各占50%的混合氮),且只加入到一侧根室(当氮加入到和PEG同侧时为水氮异区,加入到无PEG一侧时为水氮同区),测定各处理的光合、生理指标,以研究局部根区水分胁迫下氮形态与供给部位对玉米幼苗生长的影响.结果表明:同一氮形态供给下水氮同区植株的光合速率(Pn)、最大净光合速率(Pmax)、光饱和点(LSP)、CO2饱和点(CSP)、叶绿素a、b及叶绿素总含量、根系活力、氮含量和生物量高于水氮异区,光呼吸速率(Rp)、CO2补偿点(CCP)、木质部汁液脱落酸(ABA)浓度、氮利用效率、水分利用效率低于水氮异区;供混合氮和硝态氮的植株Pn、Pmax、LSP、CSP、氮含量和生物量高于供铵态氮的植株,而CCP、Rp、木质部汁液ABA浓度、氮利用效率、水分利用效率变化趋势则相反.可见,同一供氮形态下,水氮同区比水氮异区更利于植物生长,而水氮利用效率在水氮异区下较高;混合氮和硝态氮对植物生长的促进作用优于单一供给铵态氮,但铵态氮更有利于提高水氮利用效率.     

Nitrogen acquisition in four co-existing species from an upland acidic grassland

The concentration of both nitrate and ammonium nitrogen was measured in soil taken from an upland acidic (pH 4.5) grassland habitat, containing four co-existing species, Deschampsia flexuosa (L.) Trin., Festuca ovina L., Juncus squarrosus L. and Nardus stricta L. Both nitrate and ammonium nitrogen were found to be present in the soil, in similarly small quantities. The effect of both sources of nitrogen on relative growth rate was studied, and an attempt was made to determine whether nitrate or ammonium nitrogen is the immediate source of nitrogen for these plants using assays of nitrate reductase (EC 1.6.6.2) and ammonium uptake. All four species showed larger growth rates on the same concentration of ammonium nitrogen compared to nitrate nitrogen. All species showed low activities of leaf nitrate reductase, even in plants grown on 18 mol nitrate m⁻³. Ammonium uptake activity appeared to be higher in species which showed the lowest nitrate reductase activity and least response to increasing nitrate concentration in the growth medium.     

Foliar application of nitrate or ammonium as sole nitrogen supply in Ricinus communis. II. The flows of cations, chloride and abscisic acid

Following a precultivation with pedospheric nitrogen nutrition, Ricinus plants were supplied with nitrogen solely by spraying nitrate or ammonium solution onto the leaves during the experimental period. The chemical composition of tissues, xylem and phloem exudates was determined and on the basis of the previously determined nitrogen flows (Peuke et al., New Phytologist (1998), 138 , 657–687) the flows of potassium, sodium, magnesium, calcium, chloride and ABA were modelled. These data, which permit quantification of net-uptake, transport in xylem and phloem, and utilization in shoot and root, were compared with results obtained in plants with pedospherically-supplied nitrate or ammonium and data in the literature. Although the overall effects on the chemical composition of supplying ammonium to the leaves were not as pronounced as in pedospherically supplied plants, there were some typical responses of plants fed with ammonium (ammonium syndrome). In particular, in ammonium-sprayed plants uptake and transport of magnesium decreased and chloride uptake was increased compared with nitrate-sprayed plants. Furthermore, acropetal ABA transport in the xylem in ammonium-sprayed Ricinus was threefold higher than in nitrate-sprayed plants. Additionally, concentrations of anions were more or less increased in tissues, particularly in the roots, and transport fluids. The overall signal from ammonium-sprayed leaves without a direct effect of ammonium ions on uptake and transport systems in the root is discussed.     

Effects of Nitrogen Deficiency on the Absorption of Nitrate and Ammonium by Barley Plants

When young barley plants which had been supplied with nitratewere deprived of this source of N, an enhanced capacity forabsorption of either nitrate or ammonium ions developed, reachinga maximum in about 3 d under the particular experimental conditionsused. The net uptake rate of either nutrient was then approximatelythree times that in plants which had received nitrate throughout.Likewise, withholding external N from plants previously growingwith ammonium caused a 2.4-fold increase in their subsequentcapacity to absorb that ion, compared with control plants grownwith an uninterrupted ammonium supply. Accelerated nitrate uptakein N-starved plants was not accompanied by additional phosphateor sulphate absorption, but the plants had the capacity to absorbmore potassium, whether or not ammonium was also present inthe solution. Indirect evidence from analyses of root tissuesuggests that these responses to mild N-stress may depend onsome property of an N fraction which does not include nitrateor ammonium. Hordeum vulgare, barley, nitrogen, ammonium, nitrate, N-deficiency, absorption     

Uptake of Nitrate by Mutants of Arabidopsis thaliana, Disturbed in Uptake or Reduction of Nitrate

A study of nitrate and chlorate uptake by Arabidopsis thaliana was made with a wildtype and two mutant types, both mutants having been selected by resistance to high chlorate concentrations. All plants were grown on a nutrient solution with nitrate and/or ammonium as the nitrogen source. Uptake was determined from depletion in the ambient solution. Nitrate and chlorate were able to induce their own uptake mechanisms. Plants grown on ammonium nitrate showed a higher subsequent uptake rate of nitrate and chlorate than plants grown on ammonium alone. Mutant B25, which has no nitrate reductase activity, showed higher rates of nitrate and chlorate uptake than the wildtype, when both types were grown on ammonium nitrate. Therefore, the uptake of nitrate is not dependent on the presence of nitrate reductase. Nitrate has a stimulating effect on nitrate and chlorate uptake, whereas some product of nitrate and ammonium assimilation inhibits uptake of both ions by negative feedback. Mutant B 1, which was supposed to have a low chlorate uptake rate, also has disturbed uptake characteristics for nitrate.     

Pea responses to saline stress is affected by the source of nitrogen nutrition (ammonium or nitrate)

The effect of the source of nitrogen nutrition (ammonium or nitrate), onthe response of pea plants to a moderate saline stress (30 mMNaCl)was studied. Growth declined under saline stress but nitrate-fed plants wereless sensitive to salinity than ammonium-fed plants. This different sensitivitywas due mainly to a better maintenance of root growth in nitrate-fed plants.Organic nitrogen content decreased significantly in roots of ammonium-fedplants. Water relations changed slightly under saline stress leading to adecrease in stomatal conductance, which was correlated to a decline in carbonassimilation rates regardless of nitrogen source. Salinity affects the uptakeofseveral nutrients in a different way, depending on the nitrogen source. Thus,chloride was accumulated mainly in nitrate-fed plants, displacing nitrate,whereas sodium was accumulated mainly in ammonium-fed plants, especially inroots, displacing other cations such as ammonium and potassium. It is concludedthat the nitrogen source (ammonium or nitrate) is a major factor affecting pearesponses to saline stress, plants being more sensitive when ammonium is thesource used. The different sensitivity is discussed in terms of a competitionfor energy between nitrogen assimilation and sodium exclusion processes.     

Suppression of ammonium uptake by nitrogen supply and its relief during nitrogen limitation

Barley ( Hordeum vulgare , cv. Triumph), wheat ( Triticum aestivum , cv. Kleiber) and oat ( Avena sativa , cv. Tarok) were grown until day 20 in nitrate-containing solutions or in nitrogen-free solutions for periods up to 8 days immediately prior to day 20. They then were exposed for 4 h to complete, nitrate-free solutions containing 0.5 or 2.0 mM ammonium (98 atom%¹⁵N). In all 3 species in 2 experiments, net ammonium uptake was low in plants grown continuously in nitrate, and increased 3 to 4-fold with increasing nitrogen deprivation. Charge balance during net ammonium uptake was largely maintained by the sum of net potassium and net proton efflux. Variations in root ammonium concentration at the time of exposure to the ammonium solutions revealed no consistent pattern with net ammonium uptake, implying that a product of ammonium assimilation may serve as a negative effector for the uptake process. In nitrogen-replete plants, and in those deprived of nitrogen for 2 days, the amounts of endogenous ¹⁴N-ammonium recovered in the ambient ¹⁵N-ammonium solution during the 4-h uptake period were greater than the initial amounts of ¹⁴N-ammonium present in the root tissue. Significant generation of ¹⁴N-ammonium from endogenous organic nitrogen sources was thus evident in all 3 species.     

Clover and ryegrass are tolerant species to ammonium nutrition

The application of nitrification inhibitors (NIs) together with nitrogen fertilizers in grasslands is an effective alternative to reduce nitrate leaching and nitrogenous gases emissions to the atmosphere. Nevertheless, the use of NIs increases the amount of ammonium available for the plant that, due to its reported toxic effect in plants, can have a direct effect on crop production. Grassland species have traditionally suffered from intensive grazing and urea deposition and, therefore, a tolerance to ammonium nutrition could be expected in these species. Plants of Trifolium repens L. var. huia and Lolium perenne L. var. Herbus were grown under two nitrogen nutrition regimes (nitrate or ammonium) and three different nitrogen concentrations (0.5, 2.5 and 5 mmol/L). The effect of nitrogen form was determined on biomass production parameters, gas-exchange and water relations parameters as well as polyamine (PA) and ion tissue contents. Both grassland species showed tolerance to ammonium nutrition due to their capacity to adjust several metabolic processes in a species-specific way. Gas exchange measurements and biomass production (expressed as dry weight (DW)) were unaffected by the nitrogen form or dose in both species except for a decrease in root total DW in ryegrass plants grown under ammonium nutrition. Hydraulic conductance (L₀) increased in ryegrass with increasing ammonium doses but no change due to the nitrogen source was observed in water potential (Ψ_w) values. Both species, and specially ryegrass, accumulated free ammonium mainly in roots when grown under ammonium nutrition and its translocation to the shoot via xylem was also observed. A clear difference in cations and PAs pattern was observed in each species when comparing both nitrogen nutrition regimes.     

Nitrogen Use Efficiency of Crop Plants: Physiological Constraints upon Nitrogen Absorption

Current global nitrogen fertilizer use has reached approximately one hundred billion kg per annum. In many agricultural systems, a very substantial portion of this applied nitrogen fertilizer is lost from soil to groundwaters, rivers and oceans. While soil physicochemical properties play a significant part in these losses, there are several characteristic features of plant nitrogen transporter function that facilitate N losses. Nitrate and ammonium efflux from roots result in a reduction of net nitrogen uptake. As external nitrate and ammonium concentrations, respectively, are increased, particularly into the range of concentrations that are typical of agricultural soils, elevated rates of nitrate and ammonium efflux result. The rapid down-regulation of high-affinity influx as plants become nitrogen replete further reduces the root's capacity to acquire external nitrogen; only nitrogen-starved roots absorb with both high capacity and high affinity. The results of studies using molecular biology methods demonstrate that genes encoding nitrate and ammonium transporters are rapidly down-regulated when nitrogen is resupplied to nitrogen-starved plants. Provision of ammonium to roots of plants actively absorbing nitrate imposes a block on nitrate uptake, the extent of which depends on the ammonium concentration, thus further reducing the efficient utilization of soil nitrate. During the daily variation of incoming light and during periods of low incident irradiation (i.e. heavy cloud cover) the expression levels of genes encoding nitrate and ammonium transporters, and rates of nitrate and ammonium uptake, are substantially reduced. Low temperatures reduce growth and nitrogen demand, and appear to discriminate against high-affinity nitrogen influx. In sum, these several factors conspire to limit rates of plant nitrogen uptake to values that are well below capacity. These characteristics of the plant's nitrogen uptake systems facilitate nitrogen losses from soils.