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Nitrogen fertilization is common for poplar trees to improve growth and productivity. The utilization of N by poplar largely depends on fertilizer application patterns; however, the underlying regulatory hubs are not fully understood.

In this study, N utilization and potentially physiological regulations of two poplar clones XQH and BC5 were assessed through two related experiments i: five levels of N supply and ii: conventional and exponential N additions. Poplar growth leaf area and N utilization ificantly increased under fertilized compared to unfertilized conditions, whereas photosynthetic N utilization efficiency ificantly decreased under low N supplies.

Growth characteristics were better in the XQH than in the BC5 clone under the same N supplies, indicating higher N utilization efficiency. Leaf absorbed light energy, and thermal dissipation fraction was ificantly different for XQH clone between conventional and exponential N additions. Leaf concentrations of putrescine Put and acetylated Put were ificantly higher in exponential than c18 nue conventional N addition.

Photorespiration ificantly increased in leaves of XQH clone under exponential compared to conventional N addition. Our indicate that an interaction of the clone and N supply pattern ificantly occurs in poplar growth; leaf expansion and the storage N allocations are the central hubs in the regulation of poplar N utilization. Soil nitrogen N availability is one of primary factors determining plant N status.

Soil N, which exists in multi forms such as nitrate, ammonium, and amino acidsis highly mobile.

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Soil N availability is variable in natural field conditions due to biotic e. Under the background of global warming, N limitation on plant growth has been predicted to be a long-term challenge for dryland vegetation systems Hooper and Johnson, N uptake by roots is ificantly restricted under severe drought stress even if soil N is sufficient Hu and Chen, Moreover, N limitation is common in plants grown in infertile soil or good soil with short-rotation.

C18 nue of soil N availability result in inhibition of lateral root initiation, leaf development e. Deficiency of N in plant is commonly accompanied with ificant alternations of gene expressions, metabolic and physiological processes Lea et al. How to improve plant adaptation to N deficiency is a high concern in dryland agriculture. Conventional N fertilization, which delivers fertilizers at a constant rate during the growth period Hu et al.

In the past decades, excess N fertilizer was applied into global agricultural systems Good and Beatty, High N supply can produce inhibitory effects on root growth e. Appropriate fertilization managements are very crucial for crops to obtain maximum N utilization efficiency.

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Most of the plants obey the S-shaped growth pattern i. Correspondingly, plant N demand is relatively less at an initial growth stage lag phaseand rapidly increases in the log phase, and slightly decreases in the stationary phase. However inconsistent were reported in some woody species such as Eucalyptus globulus and hybrid poplar; for example, at a lower N-fertilization rate, plant morphological and nutritional traits were similar between exponentially and conventionally fertilized seedlings CloseI et al.

Plant responses to N addition are highly plastic due to the genetic-dependent such as species preference to N form and -independent such as N addition patterns factors da Silveira Pontes et al. The applicability and efficacy of exponential models for different woody species are still largely unknown.

Leaves are the major N sink of plants supplied with N fertilizers. Leaf N is primarily located in the chloroplasts in form of chloroplast proteins. During chloroplast development, polyamines PAs serve as an N source for the synthesis of chlorophyll and chloroplast proteins Sobieszczuk-Nowicka and Legocka, Polyamines are small, c18 nue molecules mainly present in chloroplasts and photosynthetic subcomplexes including thylakoid membranes, photosystem II complex and light-harvesting complex Kotzabasis et al.

Considerable evidence emphasized the importance of PAs in modulating the functionality and efficiency of photosynthetic apparatus Shu et al. PAs can stimulate leaf conversion of protochlorophyllide to chlorophyllide Beigbeder et al. Exogenous PAs have positive impacts on leaf photochemical processes in plant response to abiotic stresses Shu et al. Moreover, PAs participate in the modulation of energy production and dissipation processes of the chloroplast Ioannidis et al. Small changes of c18 nue PAs can fine tune proton motive force and ATP production in the thylakoids Ioannidis and Kotzabasis, In higher plants, free PAs are the main forms present in plant organs Chen et al.

Free PAs are implicated in the regulation of C and N metabolisms such as amino acids and tricarboxylic acid cycle. Therefore, PAs are ificantly important in leaf physiological responses to N supply.

However, to date, the information of PAs-associated plant response to exponential N addition is unavailable. Photorespiration is physiologically important in the regulation of photosynthetic carbon C and N metabolism in leaves Wang et al. In early studies e. So lowering photorespiration was operated as a potential pathway to increase leaf C fixation and plant productivity. However, increasing evidence suggests the importance of photorespiration in the regulation of N metabolism e.

Original research article

Photorespiration was implicated in plant adaptation to low N stress Tang et al. Moreover, photorespiratory pathway ificantly varies with leaf age Gjerstad, For example, the photorespiration rate is high in developing poplar leaves, whereas the value declines in the matured leaves. Leaf maturation and senescence are closely correlated with N availability Hallik et al. Photorespiratory pathway is a potentially regulatory hub in plant adaptation to N availability.

Poplar trees are short-rotation woody plants in the northern hemisphere Hart et al.

Poplar plantations have high biomass production at the cost of high water and N consumption Rockwood et al. The limitation of soil N availability is an important factor influencing the sustainability of high biomass production of the short-rotation poplar plantations.

Appropriate N applications are essential for poplars to improve plant growth and N utilization efficiency Franklin et al. Baicheng-5which are widely afforested in Northern China and have high tolerance to infertile soil, were selected as the materials.

Phenotypic variation and relationships between fatty acid concentrations and feed value of perennial ryegrass genotypes from a breeding population

It was hypothesized that 1 fertilization patterns a total amount of N supply and conventional and exponential N additions were the primary factors determining plant N status and utilization efficiency of different poplar trees; 2 leaf levels of PAs and photorespiration were implicated in poplar response to soil N availability.

Special attention was focused on storage N free amino acids, soluble proteins, PAs, and photorespiratory N allocations in leaves of the poplar clones in the growing season of post- conventional and exponential fertilization year. Cuttings of each clone were cultivated in 8. The clones were watered with tap water about ml per pot every two days. Two experiments were set up to assess 1 growth performance and leaf N utilization under five N-supply levels and 2 photosynthetic C and N c18 nue and excitation energy allocations in leaves under conventional and exponential N additions.

After plant growth for 2. N addition was conducted on July 8, July 28, and August 18 inrespectively. On the first fertilization, P 1.

Leaf N concentration, photosynthetic, and growth parameters were measured from July 8, at an interval of 20 days. Based on the of Exp. Exponential N fertilization was conducted at an interval of 10 days from June 20 to August 10,and conventional fertilization was conducted on June 20 and July 20 inrespectively.

Before sampling, non-destructive measurements [shoot height SHbasal diameter BDleaf area LAleaf photosynthesis were conducted. For the measurements of organ metabolite concentration, key enzyme activity, and fresh c18 nue leaves, stem barks, and rootssamples were separately collected in growing seasons of the fertilizing year on July 10, July 30, and post-fertilization year May 24 and June 20,respectively.

Samples collected were stored immediately in fresh boxes with ice packs. In Exp. Leaf area 5 th to 7 th leaves from the top of each plant was calculated by a c18 nue count method. Absolute water content AWC of shoots was calculated as described by Ghashghaie et al. Measurements were performed in the mornings between a. Total N was determined by an indophenol method; the detailed procedures were described by Ohyama et al. Chlorophyll fluorescence variables were calculated automatically in Handy PEA v 1. Solution concentration was determined separately using free AAs, soluble sugars, and soluble protein assay Kits Comin Biotechnology Co.

Metabolite concentration was calculated according to linear regression equations of the standard curve. Samples were diluted and derivatized as done by Gao et al. The detailed information on parameters and operations was described by Hu and Chen Elution was operated as described by Gao et al.

Mass spectrometry conditions were based on the descriptions by Jin et al.

The detailed measurements were described by Hu et al. Ltd, Suzhou, China. Enzyme activity was calculated based on the absorbance values and a standard curve.