Nitrogen Stress Affects the Turnover and Size of Nitrogen Pools Supplying Leaf Growth in a Grass

The effect of nitrogen (N) stress on the pool system supplying currently assimilated and (re)mobilized N for leaf growth of a grass was explored by dynamic ¹⁵N labeling, assessment of total and labeled N import into leaf growth zones, and compartmental analysis of the label import data. Perennial ryegrass (Lolium perenne) plants, grown with low or high levels of N fertilization, were labeled with ¹⁵NO₃⁻/¹⁴NO₃⁻ from 2 h to more than 20 d. In both treatments, the tracer time course in N imported into the growth zones fitted a two-pool model (r² > 0.99). This consisted of a “substrate pool,” which received N from current uptake and supplied the growth zone, and a recycling/mobilizing “store,” which exchanged with the substrate pool. N deficiency halved the leaf elongation rate, decreased N import into the growth zone, lengthened the delay between tracer uptake and its arrival in the growth zone (2.2 h versus 0.9 h), slowed the turnover of the substrate pool (half-life of 3.2 h versus 0.6 h), and increased its size (12.4 μg versus 5.9 μg). The store contained the equivalent of approximately 10 times (low N) and approximately five times (high N) the total daily N import into the growth zone. Its turnover agreed with that of protein turnover. Remarkably, the relative contribution of mobilization to leaf growth was large and similar (approximately 45%) in both treatments. We conclude that turnover and size of the substrate pool are related to the sink strength of the growth zone, whereas the contribution of the store is influenced by partitioning between sinks.This article examines the nitrogen (N) supply system of growing grass leaves, and it investigates how functional and kinetic properties of this system are affected by N stress. The N supply of growing leaves is a dominant target of whole-plant N metabolism. This is primarily related to the high N demand of the photosynthetic apparatus and the related metabolic machinery of new leaves (Evans, 1989; Makino and Osmond, 1991; Grindlay, 1997; Lemaire, 1997; Wright et al., 2004; Johnson et al., 2010; Maire et al., 2012). The N supply system, as defined here, is an integral part of the whole plant: it includes all N compounds that supply leaf growth. Hence, it integrates all events between the uptake of N from the environment (source), intermediate uses in other processes of plant N metabolism, and the eventual delivery to the leaf growth zone (sink; Fig. 1). N that does not ultimately serve leaf growth is not included in this system; all N that serves leaf growth is included, irrespective of its localization in the plant. Conceptually, two distinct sources supply N for leaf growth: N from current uptake and assimilation that is directly transferred to the growing leaf (“directly transferred N”) and N from turnover/redistribution of organic compounds (“mobilized N”).Open in a separate windowFigure 1.Schematic representation of N fluxes in the leaf growth zone and in the N supply system of leaf growth in a grass plant. A, Scheme of a growing leaf, with its growth zone (including zones of cell division, expansion, and maturation) and recently produced tissue (RPT). N import (I; μg h⁻¹) into the growth zone is mostly in the form of amino acids. Inside the growth zone, the nitrogenous substrate is used in new tissue construction. Then, N export (E; μg h⁻¹) is in the form of newly formed, fully expanded nitrogenous tissue (tissue-bound export with RPT) and is calculated as leaf elongation rate (L_ER; mm h⁻¹) times the lineal density of N in RPT (ρ; μg mm⁻¹): E = L_ER × ρ (Lattanzi et al., 2004). In a physiological steady state, import equals export (I = E) and the N content of the growth zone (G; μg [not shown]) is constant. Labeled N import into the growth zone (I_lab) commences shortly after labeling of the nutrient solution with ¹⁵N. The labeled N content of the growth zone (G_lab; μg) increases over time (dG_lab/dt) until it eventually reaches isotopic saturation (Fig. 2B). Similarly, the lineal density of labeled N in RPT (ρ_lab) increases until it approaches ρ. At any time, the export of labeled N in RPT (E_lab) equals the concurrent ρ_lab × L_ER. The import of labeled N is obtained as I_lab = E_lab + dG_lab/dt (Lattanzi et al., 2005) and considers the increasing label content in the growth zone during labeling. The fraction of labeled N in the import flux (f_{lab I}) is calculated as f_{lab I} = I_lab/I. The time course of f_{lab I} (Fig. 3) reflects the kinetic properties of the N supply system of leaf growth (C). B, Scheme of a vegetative grass plant (reduced to a rooted tiller with three leaves) with leaf growth zone. N import into the growth zone (I) originates from (1) N taken up from the nutrient solution that is transferred directly to the growth zone following assimilation (directly transferred N) and (2) N derived from turnover/redistribution of stores (mobilized N). The store potentially includes proteins in all mature and senescing tissue in the shoot and root of the entire plant. As xylem, phloem, and associated transfer cells/tissue provide for a vascular network that connects all parts of the plant, the mobilized N may principally originate from any plant tissue that exhibits N turnover/mobilization. The fraction of total N uptake that is allocated to the N supply system of the growth zone equals U (see model in C). The fraction of total mobilized N allocated to the growth zone equals M (see model in C). C, Compartmental model of the source-sink system supplying N to the leaf growth zone, as shown by Lattanzi et al. (2005) and used here. Newly absorbed N (U; μg h⁻¹) enters a substrate pool (Q₁); from there, the N is either imported directly into the growth zone (I) or exchanged with a store (Q₂). Q₁ integrates the steps of transport and assimilation that precede the translocation to the growth zone. Q₂ includes all proteins that supply N for leaf growth during their turnover and mobilization. The parameters of the model, including the (relative) size and turnover of pools Q₁ and Q₂, the deposition into the store (D; μg h⁻¹), and the mobilization from the store (M; μg h⁻¹), and the contribution of direct transfer relative to mobilization to the N supply of the growth zone are obtained by fitting the compartmental model to the f_{lab I} data (A) obtained in dynamic ¹⁵N labeling experiments (for details, see “Materials and Methods”). During physiological steady state, the sizes of Q₁ and Q₂ are constant, I = U, and M = D. [See online article for color version of this figure.]Amino acids are the predominant form in which N is supplied for leaf growth in grasses, and incorporation in new leaf tissue occurs mainly in the leaf growth zone (Gastal and Nelson, 1994; Amiard et al., 2004). This is a heterotrophic piece of tissue that includes the zones of cell division and elongation, is located at the base of the leaf, and is encircled by the sheath of the next older leaf (Volenec and Nelson, 1981; MacAdam et al., 1989; Schnyder et al., 1990; Kavanová et al., 2008). As most N is taken up in the form of nitrate but supplied to the growth zone in the form of amino acids, the path of directly transferred N includes a series of metabolic and transport steps. These include transfer to and loading into the xylem, xylem transport and unloading, reduction and ammonium assimilation, cycling through photorespiratory N pools, amino acid synthesis, loading into the phloem, and transport to the growth zone (Hirel and Lea, 2001; Novitskaya et al., 2002; Stitt et al., 2002; Lalonde et al., 2003; Dechorgnat et al., 2011). The time taken to pass through this sequence is unknown at present, as is the effect of N deficiency on that time. Also, it is not known how much N is contained in, and moving through, the different compartments that supply leaf growth with currently assimilated N.At the level of mature organs, mainly leaves, there is considerable knowledge about N turnover and redistribution. Much less is known about the fate of the mobilized N and its actual use in sink tissues like the leaf growth zone. The processes in mature organs are associated with the maintenance metabolism of proteins, organ senescence, and adjustments in leaf protein levels to decreasing irradiance inside growing canopies when leaves become shaded by overtopping newer ones (Evans, 1993; Vierstra, 1993; Hikosaka et al., 1994; Anten et al., 1995; Hirel et al., 2007; Jansson and Thomas, 2008; Moreau et al., 2012). N mobilization in shaded leaves supports the optimization of photosynthetic N use efficiency at plant and canopy scale (Field, 1983; Evans, 1993; Anten et al., 1995), it reduces the respiratory burden of protein maintenance costs (Dewar et al., 1998; Amthor, 2000; Cannell and Thornley, 2000), and it provides a mechanism for the conservation of the most frequently growth-limiting nutrient (Aerts, 1996). Mobilization of N involves protein turnover and net degradation (Huffaker and Peterson, 1974), redistribution in the form of amino acids (Simpson and Dalling, 1981; Simpson et al., 1983; Hörtensteiner and Feller, 2002), and (at least) some of the mobilized N is supplied to new leaf growth (Lattanzi et al., 2005).N fertilizer supply has multiple direct and indirect effects on plant N metabolism (Stitt et al., 2002; Schlüter et al., 2012). In particular, it modifies the N content of newly produced leaves, leaf longevity/senescence, and the dynamics of light distribution inside expanding canopies (Evans, 1983, 1989; Lötscher et al., 2003; Moreau et al., 2012). Thus, N fertilization influences the availability of recyclable N. At the same time, it augments the availability of directly transferable N to leaf growth. The net effect of these factors on the importance of mobilized versus directly transferred N substrate for leaf growth is not known. Also, it is unknown how N fertilization influences the functional characteristics of the N supply system, such as the size and turnover of its component pools.The assessment of the importance of directly transferred versus mobilized N for leaf growth requires studies at the sink end of the system (i.e. investigations of the N import flux into the leaf growth zone). Directly transferred N and mobilized N can be distinguished on the basis of their residence time in the plant, the time between uptake from the environment and import into the leaf growth zone: direct transfer involves a short residence time (fast transfer), whereas mobilized N resides much longer in the plant before it is delivered to the growth zone (slow transfer; De Visser et al., 1997; Lattanzi et al., 2005). Such studies require dynamic labeling of the N taken up by the plant (Schnyder and de Visser, 1999) and monitoring of the rate and isotopic composition/label content of N import into the leaf growth zone (Lattanzi et al., 2005). For grass plants in a physiological steady state, N import and the isotopic composition of the imported N are calculated from the leaf elongation rate and the lineal density of N in newly formed tissue (Fig. 1A; Lattanzi et al., 2004) and the change of tracer content in the leaf growth zone and recently produced leaf tissue over time (Lattanzi et al., 2005). Such data reveal the temporal change of the fraction of labeled N in the N import flux (f_{lab I}), which then can be used to characterize the N supply system of leaf growth via compartmental modeling. So far, there is only one study that has partially characterized this system (Lattanzi et al., 2005): this work was conducted with a C₃ grass, perennial ryegrass (Lolium perenne), and a C₄ grass, Paspalum dilatatum, growing in mixed stands and indicated that two interconnected N pools supplied the leaf growth zone in both species: a “substrate pool” (Q₁), which provided a direct route for newly absorbed and assimilated N import into the leaf growth zone (directly transferred N), and a mobilizing “store” (Q₂), which supplied N to the leaf growth zone via the substrate pool (Fig. 1C). The relative contribution of mobilization from the store was least important in the fast-growing, dominant individuals and most important in subordinate, shaded individuals. That work did not address the role of N deficiency, and the limited short-term resolution of the study (labeling intervals of 24 h or greater) precluded an analysis of the fast-moving parts of the system.Accordingly, this work addresses the following questions. How does N deficiency influence the substrate supply system of the leaf growth sink in terms of the number, size, and turnover (half-life) of its kinetically distinct pools? How does N deficiency affect the relationship between directly transferred and mobilized N for leaf growth? And what additional insight on the compartmental structure of the supply system is obtained when the short-term resolution of the analysis is increased by 1 order of magnitude? The work was performed with vegetative plants of perennial ryegrass grown in constant conditions with either a low (1.0 mm; termed low N) or high (7.5 mm; high N) nitrate concentration in the nutrient solution. In both treatments, a large number of plants were dynamically labeled with ¹⁵N over a wide range of time intervals (2 h to more than 20 d). The import of total N and ¹⁵N tracer into growth zones was estimated at the end of each labeling interval. Tracer data were analyzed with compartmental models following principles detailed by Lattanzi et al. (2005, 2012) and Lehmeier et al. (2008) to address the specific questions. Previous articles reported on root and shoot respiration (Lehmeier et al., 2010) and cell division and expansion in leaf growth zones (Kavanová et al., 2008) in the same experiment.     

Effects of Rootstocks and Interstocks on the Xylem Sap Composition in Apple Trees: Effects on Nitrogen, Phosphorus, and Potassium Content

This paper records the concentration of N, P, and K in sap whichexuded from the cut stumps of composite orchard apple trees.The trees were either single-worked on different rootstocksor incorporated various interstocks. While the volume of sap exuded per unit area of cut surfacedid not appear to vary with rootstock or interstock, exudatefrom the dwarfing rootstocks or interstocks contained depletedlevels of nutrients. A possible mechanism for rootstock and interstock effects isdiscussed in the light of these results.     

Effect of Dwarfing Interstocks on Xylem Sap Composition in Apple Trees: Effect on Nitrogen, Potassium, Phosphorus, Calcium and Magnesium Content

This paper concerns nutrient concentrations in xylem sap collectedfrom above and below M.8, M.9, M.26 and M.7 dwarfing interstocks.The sap flowing from above an interstock had lower concentrationsof nutrients than that from the rootstock levels and these differencesincreased with the dwarfing effect of the interstock. There were indications that nutrient concentrations were enhancedbelow interstocks but were reduced in the sap flowing from theinterstocks into the scions. Generally these reductions weregreater the more dwarfing the effect of the interstock. Analyses of sap from above, mid-way along and below interstocksindicated that the changes in content were produced in, or closeto, the graft union between scion and interstock.     

Absorption of Nitrogen, Phosphorus, and Potassium from Nutrient Sprays by Leaves

Barley, Brussels sprout, French bean, tomato, and sugar-beetplants grown in soil in pots and sprayed, usually daily, forseveral weeks, with nutrient solutions containing nitrogen,phosphorus, potassium, and a spreader, with precautions to preventthe spray solution falling on the soil, had higher nutrientcontents and dry weights than control plants sprayed with waterand spreader only. Increase in nutrient content occurred withhigh or low levels of nutrient supply to the roots and was approximatelyproportional to the concentration of spray and to the frequencyof spraying. The nitrogen content of sugar-beet plants was increased equallyby spraying with solutions supplying ammonium sulphate, calciumnitrate, or urea in equivalent concentrations. Nutrient uptake from solutions sprayed on leaves influenceduptake by the roots so that the additional amounts of nutrientcontained in sprayed plants may be greater or smaller than theamount absorbed from the spray by the leaves.     

Influence of Nitrogen Supply and Water Stress on Growth and Nitrogen, Phosphorus, Potassium and Calcium Contents in Pearl Millet

Influence of supra-optimal concentrations of N on growth and accumulation of N, K, P and Ca in the shoots and roots in Pennisetum glaucum (L.) R.Br. under water stress was assessed in a pot experiment under glasshouse conditions. Thirty four-day-old plants of two lines, ICMV94133 and WCA-78, were subjected to 224, 336, or 448 mg(N) kg^–1(soil) and soil moisture 100 or 30 % of field capacity for 30 d. Increasing soil N supply decreased growth of both lines under water deficit. Nitrogen content in the shoots of both lines was not affected by supra-optimal levels of N or different watering regimes, but in contrast, the root N content was increased consistently in WCA-78 with increase in soil N content. Shoot P content increased considerably in WCA-78 at the two higher N contents, but it was significantly lower at drought stress than at well-watered treatment. In contrast, shoot or root P content in ICMV94133 did not differ under both watering regimes. Potassium content in the shoots of WCA-78 was considerably increased at the two higher N contents under drought conditions. Root K content was increased in WCA-78 at the highest N content under well-watered conditions, whereas the reverse was true in ICMV94133. Calcium content in the shoots of ICMV94133 was higher under drought stress compared with that at well-watered conditions, but such pattern was not observed in WCA-78. However, root Ca content increased in both lines with increase in N supply.     

Carbon,Nitrogen, Phosphorus,and Potassium Stoichiometry in an Ombrotrophic Peatland Reflects Plant Functional Type

Ombrotrophic bog peatlands are nutrient-deficient systems and important carbon (C) sinks yet the stoichiometry of nitrogen (N), phosphorus (P) and potassium (K), essential for plant growth and decomposition, has rarely been studied. We investigated the seasonal variation in C, N, P, and K concentrations and their stoichiometric ratios in photosynthetically active tissues of 14 species belonging to five plant functional types (PFTs) (mosses, deciduous trees/shrubs, evergreen shrubs, graminoids, and forb) at Mer Bleue bog, an ombrotrophic peatland in eastern Ontario, Canada. Although we observed variations in stoichiometry among PFTs at peak growing season, there was convergence of C:N:P:K to an average mass ratio of 445:14:1:9, indicating N and P co-limitation. Nitrogen, P, and K concentrations and stoichiometric ratios showed little seasonal variation in mosses, evergreens, and graminoids, but in forb and deciduous species were the largest in spring and decreased throughout the growing season. Variations in nutrient concentrations and stoichiometric ratios among PFTs were greater than seasonal variation within PFTs. Plants exhibit N and P co-limitation and adapt to extremely low nutrient availability by maintaining small nutrient concentrations in photosynthetically active tissues, especially for evergreen shrubs and Sphagnum mosses. Despite strong seasonal variations in nutrient availabilities, few species show strong seasonal variation in nutrient concentrations, suggesting a strong stoichiometric homeostasis at Mer Bleue bog.     

Variation in Yield Gap Induced by Nitrogen,Phosphorus and Potassium Fertilizer in North China Plain

A field experiment was conducted under a wheat-maize rotation system from 1990 to 2006 in North China Plain (NCP) to determine the effects of N, P and K on yield and yield gap. There were five treatments: NPK, PK, NK, NP and a control. Average wheat and maize yields were the highest in the NPK treatment, followed by those in the NP plots among all treatments. For wheat and maize yield, a significant increasing trend over time was found in the NPK-treated plots and a decreasing trend in the NK-treated plots. In the absence of N or P, wheat and maize yields were significantly lower than those in the NPK treatment. For both crops, the increasing rate of the yield gap was the highest in the P omission plots, i.e., 189.1 kg ha⁻¹ yr⁻¹ for wheat and 560.6 kg ha⁻¹ yr⁻¹ for maize. The cumulative omission of P fertilizer induced a deficit in the soil available N and extractable P concentrations for maize. The P fertilizer was more pivotal in long-term wheat and maize growth and soil fertility conservation in NCP, although the N fertilizer input was important for both crops growth. The crop response to K fertilizers was much lower than that to N or P fertilizers, but for maize, the cumulative omission of K fertilizer decreased the yield by 26% and increased the yield gap at a rate of 322.7 kg ha⁻¹ yr⁻¹. The soil indigenous K supply was not sufficiently high to meet maize K requirement over a long period. The proper application of K fertilizers is necessary for maize production in the region. Thus, the appropriate application of N and P fertilizers for the growth of both crops, while regularly combining K fertilizers for maize growth, is absolutely necessary for sustainable crop production in the NCP.     

Interactions of Nitrogen, Phosphorus, and Potassium supplied in Leaf Sprays or in Fertilizer added to the Soil

When sugar-beet plants grown in pots were sprayed daily withnutrient solutions supplying nitrogen, phosphorus, and potassiumseparately or in all combinations, with precautions to preventspray falling on the soil in which the plants were grown, allthree nutrients were absorbed through the leaves. In one experimentnitrogen and potassium, and in another only nitrogen, causedincreases in plant dry weight and leaf area. Swedes absorbedphosphorus from leaf sprays and from fertilizer applied to thesoil, but only the fertilizer caused an increase in dry weight. Absorption of any of the nutrients tested from a spray containingmore than one nutrient was unaffected by the presence of othersin the spray, but spraying with nitrogen-containing solutionsincreased the absorption of phosphorus and potassium from thesoil, and potassium in sprays increased the uptake of phosphorusfrom the soil. Nitrogenous fertilizer applied to the soil increased the leafarea of sugar-beet plants, and hence it also increased the amountsof nitrogen, phosphorus, and potassium deposited on the leaveswhen they were sprayed with solutions of these nutrients, andthe amounts absorbed from the spray into the plants. Phosphaticfertilizer had no effect on uptake from leaf sprays. Potassicfertilizer did not affect leaf area or the estimated volumeof spray solution retained on the leaves, but it appeared toreduce uptake of potassium from the spray. Dry weight per plant was increased by all three nutrients infertilizer, and sugar yield of the roots was increased by nitrogenand potassium in fertilizer, and by nitrogen in spray. Applicationof a nutrient in leaf spray reduced the responses in dry weightand sugar yield to the same nutrient applied in fertilizer tothe soil. Less nitrogen, but more phosphorus, was taken up from the leafsprays than from fertilizer. Nutrients from sprays producedsmaller increases in total dry weight and in dry weight perunit of absorbed nutrient than the same nutrient from fertilizer. The apparent percentage recovery of nitrogen applied in spray,based on estimates of the volumes of solution retained on theleaves, was unaffected by fertilizer treatment, that of phosphoruswas increased by nitrogen fertilizer, and that of potassiumwas increased by nitrogen fertilizer and reduced by potassiumfertilizer. The volume of spray solution held on the leaveswas probably overestimated, so that the highest apparent recovery,about 60 per cent., may represent an almost complete true recovery,because only trivial amounts of the nutrients that had beenapplied in spray remained on the leaf surface to be removedby washing before harvest. Lower apparent recoveries may bedue to reduced uptake from the soil of the nutrient suppliedin spray.     

Nitrogen, Phosphorus, and Potassium Nutrition of the Edaphic Ecotypes in Cynodon dactylon (L.) Pers.

A differential response to nitrogen, phosphorus, and potassiumwas observed for the three edaphic ecotypes in Cynodon dactylon(L.) Pers. naturally occurring at Malla, Shiwalik, and Chandigarh.The soils of these sites differ markedly with respect to availablenutrients. The better growth of the Malla population comparedto the Chandigarh population in low potassium and low nitrogenmedia could be related to the low availability of these nutrientsin the calcareous Malla soil. The Shiwalik population tendedto show an intermediate behaviour. Whilst the Malla populationwas susceptible to ‘phosphate toxicity’, the growthyield of the Shiwalik and the Chandigarh populations improvedwith increase in phosphorus in the medium. The three populationsalso differed from each other with respect to the uptake ofvarious nutrients. These results could be correlated with thenatural habitat conditions in which the three ecotypes occur.It was concluded that besides calcium which was shown to bepartly responsible for the restriction of the three ecotypesto their respective habitats, the availability of nitrogen,potassium, and also phosphorus may play an important role.     

Identification of Nitrogen,Phosphorus, and Potassium Deficiencies in Rice Based on Static Scanning Technology and Hierarchical Identification Method

Establishing an accurate, fast, and operable method for diagnosing crop nutrition is very important for crop nutrient management. In this study, static scanning technology was used to collect images of a rice sample''s fully expanded top three leaves and corresponding sheathes. From these images, 32 spectral and shape characteristic parameters were extracted using an RGB mean value function and using the Regionprops function in MATLAB. Hierarchical identification was used to identify NPK deficiencies. First, the normal samples and non-normal (NPK deficiencies) samples were identified. Then, N deficiency and PK deficiencies were identified. Finally, P deficiency and K deficiency were identified. In the identification of every hierarchy, SVFS was used to select the optimal characteristic set for different deficiencies in a targeted manner, and Fisher discriminant analysis was used to build the diagnosis model. In the first hierarchy, the selected characteristics were the leaf sheath R, leaf sheath G, leaf sheath B, leaf sheath length, leaf tip R, leaf tip G, leaf area and leaf G. In the second hierarchy, the selected characteristics were the leaf sheath G, leaf sheath B, white region of the leaf sheath, leaf B, and leaf G. In the third hierarchy the selected characteristics were the leaf G, leaf sheath length, leaf area/leaf length, leaf tip G, difference between the 2^nd and 3^rd leaf lengths, leaf sheath G, and leaf lightness. The results showed that the overall identification accuracies of NPK deficiencies were 86.15, 87.69, 90.00 and 89.23% for the four growth stages. Data from multiple years were used for validation, and the identification accuracies were 83.08, 83.08, 89.23 and 90.77%.     

Loss of Nitrogen,Phosphorus, and Potassium Through Crop Harvests in Agroecosystems of Qianjiang,Hubei Province,PR China

The level of chemical fertilization in China is considerably higher than that normally applied in Western countries. For example, in the last decade the use of N-P-K chemical fertilizers increased, within the province of our study, from about 300 kg/ha in 1980 to about 850 kg/ha in 1990. Data on total amounts on nitrogen, phosphorus, and potassium that are removed with the harvest are necessary in determining and optimizing the amount of chemical fertilizers to be applied to different crops and in different cropping patterns. In this article we report the results of analyses of N, P, and K content of soils and different parts of different plants (soybean, cotton, rice, peanut, radish, Chinese cabbage, sesame, pepper, eggplant, and ambary) typically grown in rural areas of Southern China and provide an estimate of the balance of these nutrients under the five cropping systems that are dominant in the studied area.     

Biomass,Carbon, and Nitrogen Pools in Mexican Tropical Dry Forest Landscapes

Tropical dry forest is the most widely distributed land-cover type in the tropics. As the rate of land-use/land-cover change from forest to pasture or agriculture accelerates worldwide, it is becoming increasingly important to quantify the ecosystem biomass and carbon (C) and nitrogen (N) pools of both intact forests and converted sites. In the central coastal region of México, we sampled total aboveground biomass (TAGB), and the N and C pools of two floodplain forests, three upland dry forests, and four pastures converted from dry forest. We also sampled belowground biomass and soil C and N pools in two sites of each land-cover type. The TAGB of floodplain forests was as high as 416 Mg ha^–1, whereas the TAGB of the dry forest ranged from 94 to 126 Mg ha^–1. The TAGB of pastures derived from dry forest ranged from 20 to 34 Mg ha^–1. Dead wood (standing and downed combined) comprised 27%–29% of the TABG of dry forest but only about 10% in floodplain forest. Root biomass averaged 32.0 Mg ha^–1 in floodplain forest, 17.1 Mg ha^–1 in dry forest, and 5.8 Mg ha^–1 in pasture. Although total root biomass was similar between sites within land-cover types, root distribution varied by depth and by size class. The highest proportion of root biomass occurred in the top 20 cm of soil in all sites. Total aboveground and root C pools, respectively, were 12 and 2.2 Mg ha^–1 in pasture and reached 180 and 12.9 Mg ha^–1 in floodplain forest. Total aboveground and root pools, respectively, were 149 and 47 kg ha^–1 in pasture and reached 2623 and 264 kg ha^–1 in floodplain forest. Soil organic C pools were greater in pastures than in dry forest, but soil N pools were similar when calculated for the same soil depths. Total ecosystem C pools were 306. The Mg ha^–1 in floodplain forest, 141 Mg ha^–1 in dry forest, and 124 Mg ha^–1 in pasture. Soil C comprised 37%–90% of the total ecosystem C, whereas soil N comprised 85%–98% of the total. The N pools lack of a consistent decrease in soil pools caused by land-use change suggests that C and N losses result from the burning of aboveground biomass. We estimate that in México, dry forest landscapes store approximately 2.3 Pg C, which is about equal to the C stored by the evergreen forests of that country (approximately 2.4 Pg C). Potential C emissions to the atmosphere from the burning of biomass in the dry tropical landscapes of México may amount to 708 Tg C, as compared with 569 Tg C from evergreen forests.     

Oxidative Stress and Antioxidant Responses in Young Leaves of Mulberry Plants Grown Under Nitrogen, Phosphorus or Potassium Deficiency

The aim of this study was to associate the generation of reactive oxygen species （ROS） with Induced antloxidant responses and disturbed cellular redox environment in the nitrogen-（N）, phosphorus-（P）, or potassium-（K） deftcient mulberry （Morus alba L. var. Kanva-2） plants. The indicators of oxidative stress and cellular redox environment and antioxldant defense-related parameters were analyzed. Oeficlency of N, P or K suppressed growth, accelerated senescence, and decreased concentrations of chloroplastic pigments and glutathione. Lipid peroxidation and activities of superoxide dismutase, ascorbate peroxidase and glutathione reductase were also increased in these N, P, or K deprived plants. Concentration of hydrogen peroxide Increased in plants deficient in N or P. Oeficlency of N or P particularly altered the cellular redox environment as indicated by changes in the redox couples, namely ascorbic acid/total ascorbate decreased in P-, glutathione sulfydryl/total glutathione decreased in N-, and Increased in P-deficient plants. Activity staining of native gels for superoxide dismutase revealed Increased activity as Indicated by Increased intensity of bands, and induction of few new isoforms in P- and K-deficient plants. Oifferences in the patterns of superoxide dismutase isoforms and redox status （ascorbic acid/total ascorbate and glutathlone sulfydryl/total glutathione） Indicate that N-, P-, or K-deficiency altered antioxidant responses to varying extents in mulberry plants.     

黄芩干物质累积和氮磷钾吸收、分配规律研究

为了探讨黄芩干物质累积和氮、磷、钾吸收与分配的特点及两者间的相互关系,通过田间试验和采样分析,研究了黄芩不同生育期植株的干物质和氮、磷、钾累积量.结果表明,黄芩干物质的累积量随生育进程不断地增加,出苗后52～85 d干物质累积量占总累积量的61.62％.在整个生育期,黄芩对K2O的吸收累积量最大,N次之,P2O5最小,N、P2O5、K2O吸收比例约为2.8∶1.0∶2.9,并且黄芩地上部氮磷钾的累积量大于根部,不同生育期,根部N、P2O5、K2O的累积比例呈现增加—降低—增加的趋势.黄芩对氮磷钾的积累量与干物质积累量呈极显著正相关关系.在供试的土壤和施肥条件下,每生产100 kg的黄芩根需要从土壤和肥料中吸收6.34 kg的N,2.60 kg的P2O5,7.02 kg的K2O.     

Short-Term Response of Switchgrass to Nitrogen,Phosphorus, and Potassium on Semiarid Sandy Wasteland Managed for Biofuel Feedstock

It is important to understand switchgrass (Panicum virgatum L.) productivity with relation to diverse nutrient deficiency conditions in order to optimize continuous biomass production in marginal lands. This study was conducted on a wasteland sandy soil (Aridosol) to assess biomass yield, nutrient uptake and nitrogen (N) recovery of switchgrass, and soil nitrate-N (NO₃^?-N) accumulation responses to N (120 kg N ha^?1), phosphorus (P, 100 kg P₂O₅ ha^?1), and potassium (K, 45 kg K₂O ha^?1) applications during 2015 and 2016 in Inner Mongolia, China. The experiment layout was a randomized complete block design with fertilizer mixture treatments of N, P, and K (NPK), P and K (PK), N and K (NK), N and P (NP), and a control with no fertilizer input (CK). Plant height and stem diameter remained unaffected by the different fertilizer treatments. Biomass yield with the NPK treatment in 2015 was 8.9 Mg ha^?1 and in 2016 it was 7.3 Mg ha^?1. In 2015, compared with the NPK treatment, a significant yield reduction of 33.7% was found with PK, 22.5% with NK, 28.1% with NP, and 40.5% with CK; however, in 2016, yield declined significantly only with CK compared to the rest of the fertilizer treatments, for which yields were statistically similar. Plant N content was reduced for the treatment PK (i.e. N omission); conversely, plant P and K content remained unaffected with P and K omission treatments. Plant nutrient uptake, particularly of N and K, was severely decreased by the nutrient omission treatments when averaged across 2 years. Apparent N recovery (ANR; quantity of N uptake per unit of N applied) was reduced for the NP and NK treatments, which led to an increase in soil NO₃^?-N accumulation in the top 0–20 cm layer, compared with the NPK treatment. However, ANR was the highest (37.2% in 2015) with the NPK treatment, which also reduced soil NO₃^?-N accumulation. A balanced N, P, and K fertilizer management approach is suggested to sustain switchgrass yield and stand persistence on semiarid, marginal, sandy wasteland.     

氮、磷、钾营养对膜荚黄芪幼苗根系活力和游蓠氨基酸含量的影响

运用水培试验法研究不同营养水平对黄芪幼苗根系活力和游离氨基酸组成及含量的影响。结果表明：缺素显著降低黄芪根系活力,不同营养处理游离氨基酸含量差异显著,游离氨基酸总量的变化规律为叶片〉根,各处理游离氨基酸总量为-K〉-P〉NPK〉-N。全素处理与缺素处理相比．能提高根系活力、协调根冠比。黄芪幼苗通过提高体内游离氨基酸含量以增强对营养胁迫逆境的适应能力。     

莲藕干物质和氮磷钾养分的累积与分配研究

连续2年采用盆栽试验研究了莲藕（Nelumbo nucifera Gaertn）干物质和氮磷钾养分的累积与分配规律。结果表明：莲藕苗期以叶片生长并积累光合产物为主,膨大根状茎成型后,叶片、叶柄和根状茎中的干物质不断运输并贮存到膨大根状茎中,以产量形成为主,干物质累积总量增长呈＂慢-快-稳定＂的变化趋势;氮磷钾累积量与干物质累积量变化趋势一致,并与之呈极显著正相关,莲藕氮磷钾养分累积总量之比为1∶0.12∶1.31。移栽后97-160 d是莲藕产量形成的关键时期,不仅叶片、叶柄和根状茎中的氮磷钾随同干物质运输并贮存到膨大根状茎中,根系还从土壤中吸收更多的氮磷钾直接运输并贮存到膨大根状茎中,后者分别占同期氮磷钾累积量的69.8%、79.2%和75.0%。160 d膨大根状茎中干物质、氮、磷和钾累积量分别平均占植株总累积量的81.1%、85.2%、88.8%和80.2%。     

氮、磷、钾营养对膜荚黄芪幼苗根系活力和游离氨基酸含量的影响

运用水培试验法研究不同营养水平对黄芪幼苗根系活力和游离氨基酸组成及含量的影响。结果表明:缺素显著降低黄芪根系活力,不同营养处理游离氨基酸含量差异显著,游离氨基酸总量的变化规律为叶片>根,各处理游离氨基酸总量为-K>-P>NPK>-N。全素处理与缺素处理相比,能提高根系活力、协调根冠比。黄芪幼苗通过提高体内游离氨基酸含量以增强对营养胁迫逆境的适应能力。     

不同滴灌定额对春播裸燕麦氮磷钾质量分数及产量性状的影响

在大田条件下,采用随机区组设计,研究不同灌溉定额对春播裸燕麦不同生育时期氮磷钾质量分数以及产量性状的影响。结果表明, 裸燕麦高灌溉定额处理不同生育时期茎、叶和穗中氮磷钾的质量分数显著高于低灌溉定额(P<0.05)。滴灌灌溉定额为180 mm（W4处理）时不同生育时期各器官氮磷钾质量分数最高,其他处理由高到低依次为：W5、CK、W3、W2、W1;滴灌灌溉定额为60 mm（W1处理）不同生育时期各器官氮磷钾质量分数最低。滴灌灌溉定额为180 mm（W4处理）时籽粒产量最高（3 249.7 kg/hm）,比传统灌溉（CK处理）籽粒产量提高5.98%,比最低灌溉定额（W1处理）提高31.83%。说明适宜的滴灌灌溉定额一定程度上可以促进裸燕麦不同生育时期茎、叶和穗对氮、磷、钾的吸收,使得各器官养分的积累量增加,促使后期有更多养分向穗部转移,从而有利于燕麦籽粒产量的提高。     

Effects of Mechanized,Deep Application of Slow-Release Fertilizer on Yield and Nitrogen,Phosphorus, and Potassium Utilization of Direct-Seeded Rice

Compared with the standard method of manual fertilizer broadcasting (MFB), mechanized hill-drilling direct-seeding with deep application of slow-release nitrogen fertilizer (MHDDF) is an efficient method to integrate both fertilization and seeding. However, there are few studies that combine the use of slow-release fertilizer with MHDDF. We sought to explore the combined effect of MHDDF with slow-release fertilizer on rice yield and nitrogen, phosphorus, and potassium utilization, compared to MFB. We compared three different MHDDF methods (D30: 450 kg ha^?1, D40: 600 kg ha^?1, D50: 750 kg ha^?1), with one MFB method (B50: 750 kg ha^?1), and one control (CK: 0 kg ha^?1). We found that the yield of all MHDDF method was higher than that of both the MFB method. Yield was the highest in the D50 treatment and was 14.14–46.03% higher than that in B50 treatment. Biomass accumulation, nutrient accumulation, and nutrient use efficiency were similarly higher in MHDDF method than both MFB and CK. Compared to B50, the D50 treatment increased nitrogen recovery efficiency by 170.53–231.50%, phosphorus recovery efficiency by 480.00–724.25%, and potassium recovery efficiency by 201.55–169.59%. Overall, we found that combining MHDDF with slow-release fertilizer was an effective method to increase rice yield and nutrient use efficiency compared with MFB.