Effect of nitrogen and phosphorus addition on leaf nutrient concentrations and nutrient resorption efficiency of two dominant alpine grass species

Nitrogen(N)and phosphorus(P)are two essential nutrients that determine plant growth and many nutrient cycling processes.Increasing N and P deposition is an important driver of ecosystem changes.However,in contrast to numerous studies about the impacts of nutrient addition on forests and temperate grasslands,how plant foliar stoichiometry and nutrient resorption respond to N and P addition in alpine grasslands is poorly understood.Therefore,we conducted an N and P addition experiment(involving control,N addition,P addition,and N+P addition)in an alpine grassland on Kunlun Mountains(Xinjiang Uygur Autonomous Region,China)in 2016 and 2017 to investigate the changes in leaf nutrient concentrations(i.e.,leaf N,Leaf P,and leaf N:P ratio)and nutrient resorption efficiency of Seriphidium rhodanthum and Stipa capillata,which are dominant species in this grassland.Results showed that N addition has significant effects on soil inorganic N(NO_(3)^(-)-N and NH_(4)^(+)-N)and leaf N of both species in the study periods.Compared with green leaves,leaf nutrient concentrations and nutrient resorption efficiency in senesced leaves of S.rhodanthum was more sensitive to N addition,whereas N addition influenced leaf N and leaf N:P ratio in green and senesced leaves of S.capillata.N addition did not influence N resorption efficiency of the two species.P addition and N+P addition significantly improved leaf P and had a negative effect on P resorption efficiency of the two species in the study period.These influences on plants can be explained by increasing P availability.The present results illustrated that the two species are more sensitive to P addition than N addition,which implies that P is the major limiting factor in the studied alpine grassland ecosystem.In addition,an interactive effect of N+P addition was only discernable with respect to soil availability,but did not affect plants.Therefore,exploring how nutrient characteristics and resorption response to N and P addition in the alpine grassland is important to understand nutrient use strategy of plants in terrestrial ecosystems.

Leaf nutrient dynamics and nutrient resorption:a comparison between larch plantations and adjacent secondary forests in Northeast China

Aims Conversion of secondary forests to pure larch plantations is a common management practice driven by the increasing demand for timber production in Northeast China,resulting in a reduction in soil nutrient availability after a certain number of years following conversion.Nutrient resorption prior to leaf senescence was related to soil fertility,an important nutrient conservation strategy for plants,being especially significant in nutrient-poor habitats.However,the seasonal dynamics of leaf nutrients and nutrient resorption in response to secondary forest conversion to larch plantations is not well understood.Methods A comparative experiment between larch plantations(Larix spp.)and adjacent secondary forests(dominant tree species including Quercus mongolica,Acer mono,Juglans mandshurica and Fraxinus rhynchophylla)was conducted.We examined the variations in leaf nutrient(macronutrients:N,P,K,Ca and Mg;micronutrients:Cu and Zn)concentrations of these tree species during the growing season from May to October in 2013.Nutrient resorption efficiency and proficiency were compared between Larix spp.and the broadleaved species in the secondary forests.Important Findings Results show that the seasonal variation of nutrient concentrations in leaves generally exhibited two trends,one was a downward trend for N,P,K,Cu and Zn,and another was an upward trend for Ca and Mg.The variations in foliar nutrient concentrations were mainly controlled by the developmental stage of leaves rather than by tree species.Resorption of the observed seven elements varied among the five tree species during leaf senescence.Nutrient resorption efficiency varied 6–75%of N,P,K,Mg,Cu and Zn,while Ca was not retranslocated in the senescing leaves of all species,and Mg was not retranslocated in Larix spp.Generally,Larix spp.tended to be more efficient and proficient(higher than 6–30%and 2–271%of nutrient resorption efficiency and resorption proficiency,respectively)in resorbing nutrients than the broadleaved species in the secondary forests,indicating that larch plantations had higher leaf nutrient resorption and thus nutrient use efficiency.Compared with Larix spp.,more nutrients would remain in the leaf litter of the secondary forests,indicating an advantage of secondary forests in sustaining soil fertility.In contrast,the larch plantation would reuse internal nutrients rather than lose nutrients with litter fall and thus produce a positive feedback to soil nutrient availability.In summary,our results suggest that conversion from secondary forests to pure larch plantations would alter nutrient cycling through a plantmediated pathway.

N limitation increases along a temperate forest succession:evidences from leaf stoichiometry and nutrient resorption

Forest productivity and carbon(C) sequestration largely depend on soil N and P availability.To date,however,the temporal variation of nutrient limitation along forest succession is still under debate.Leaf stoichiometry and nutrient resorption are important indicators for predicting nutrient limitation of plant growth.Here,we measured nitrogen(N)and phosphorus(P)concentrations in green leaves and leaf liter for all woody species at four stages of temperate forest succession,and analyzed how abiotic and biotic factors affect leaf stoichiometry and nutrient resorption along forest succession.At the individual scale,leaf N and P concentrations had a significant increase at the end of the succession,while no change in leaf N:P ratio was detected.Nitrogen resorption efficiency(NRE)increased significantly with succession,but P resorption efficiency(PRE)first increased and then decreased.Significant increases in NRE:PRE ratios only occurred at the end of the succession.Moreover,plant N cycling was less responsive to soil nutrient than P cycling.At the community scale,we found that leaf N and P concentrations first decreased and then increased along forest succession,which were mainly affected by Shannon-Wiener index and species richness.Leaf N:P ratio significantly varied with succession and was mainly determined by community-weighted mean diameter at breast height(DBH).NRE increased and was significantly influenced by species richness and DBH,while PRE was relatively stable along forest succession.Thus,the NRE:PRE ratios significantly increased,indicating that N limitation is exacerbated with the temperate forest succession.These results might reflect the intense interspecific competition for limiting resource in a higher biodiversity community.In conclusion,our findings highlight the importance of biotic factors in driving forest ecosystem nutrient cycling and provide valuable information for sustainable fertilizer management practices in China's temperate and boreal forests.

Nutrient resorption and stoichiometric responses of poplar(Populus deltoids)plantations to N addition in a coastal region of eastern China

Aims Leaf nutrient resorption is sensitive to changes in soil nutrients.However,the effects of N deposition on nutrient resorption efficiency(NuRE)in plant macro-nutrients remain unclear.Poplar(Populus deltoids)is one of the most extensively cultivated hardwood species worldwide.We explored general patterns and dominant drivers of NuRE and stoichiometry of poplar plantations in response to N addition.Methods We conducted a 4-year N-addition experiment to explore NuRE and stoichiometric responses to N addition in two poplar(P.deltoids)plantations(8-and 12-year-old stands)in a coastal region of eastern China.We measured soil and foliar(green and senesced leaves)concentrations of nitrogen(N),phosphorus(P),potassium(K),calcium(Ca)and magnesium(Mg)for a series of N addition treatments including N_(0)(0 kg N ha^(−1)yr^(−1)),N_(1)(50 kg N ha^(−1)yr^(−1)),N_(2)(100 kg N ha^(−1)yr^(−1)),N_(3)(150 kg N ha^(−1)yr^(−1))and N_(4)(300 kg N ha^(−1)yr^(−1)).Important Findings Consistent for(both)8-and 12-year-old stands,N addition did not affect the NuRE and stoichiometry(with the exception of CaRE and CaRE:MgRE ratio).N resorption efficiency–P resorption efficiency(NRE–PRE)scaling slopes were consistently less than 1.0 under N addition.These results suggest that NRE generally decouples from PRE within each N treatment.Moreover,these results point to robust control of green leaf nutritional status on nutrient resorption processes as indicated by the positive relationships between NuRE and green leaf nutrient concentrations.Our findings provided a direct evidence that growth in 12-year-old poplar plantations was N-limited in the coastal region of eastern China.

Experimental evidence for weakened tree nutrient use and resorption efficiencies under severe drought in a subtropical monsoon forest

Aims The functions of global forests are threatened by the increasing frequency of severe drought.Due to drought inducing reductions in soil nutrient availability,efficiencies of nutrient use and resorption of trees become crucial for forest functions and biogeochemical cycles.However,understanding the dynamics of responses of foliar nutrient use and resorption efficiencies to drought,especially in tropical or subtropical forests,is still limited.Our goal was to detect whether and how the importance of leaf nutrient use and resorption changes across different species in the hot and wet forests when suffering drought stress in different months.Methods Based on a 70%throughfall exclusion experiment in a subtropical forest,we collected green and senesced leaves of Schima superba and Lithocarpus glaber in different months from October 2016 to May 2019,to estimate the effects of drought on leaf nitrogen(N)and phosphorus(P)use and resorption efficiencies(i.e.NUE and PUE,NRE and PRE).Important Findings The effects of drought on nutrient use and resorption efficiencies varied between species and months.Based on a 2-year observation,drought had no effect on S.superba,but significantly decreased NUE,NRE and PRE of L.glaber by 3.4%,20.2%and 7.1%,respectively.Furthermore,the negative drought effects were aggravated by the natural summer drying in 2017.As a result,NUE and PUE of L.glaber were significantly depressed by 17.2%and 58.1%,while NRE and PRE were significantly reduced by 56.5%and 53.8%in August 2017.Moreover,the responses of NRE,PRE and NUE to drought were related with soil moisture(SM)for L.glaber,and when SM decreased to a threshold near 9 v/v%,drought effects were shifted from unresponsive to negative.Our results highlight a species-specific threshold response of nutrient use under drought in a subtropical forest.

Seasonal changes of bioelements in litter and their potential return to green leaves in five species of tropical dry deciduous forest, western India

The Litter nutrient concentrations of N, P, Ca, Mg, K and Na in mature leaves, twigs and reproductive parts and their relationship between senescent and young leaves were investigated in five forest species： Acacia nilotica, Acacia leucophloea. Tectona grandis, Miliusa tomentosa and Butea monosperma in Indian tropical dry deciduous forest in Western India.. Total dry matter of plant species was recorded and analyzed for N, P, Ca, Mg, K and Na. A. nilotica had the highest concentrations of N in leaf, while ,4. leucophloea had the highest concentrations of Ca and Mg in leaf. The highest concentrations of P in leaf were found in A. nilotica, A. leucophloea and B. monosperma where as lowest in T. grandis and M. tomentosa. No significant differences in K and Na were registered among the species. A marked seasonal variability was ob- served in the concentrations of N, P and K, except for Ca and Mg. Potas- sium is the single element that undergoes leaching and mobilization in all species. Resorped N and P can be used for the production of fresh leaf in the following annual cycle. Nutrient resorption and retranslocation from senescent leaves and litter supports the production of new foliage and increase the fertility of soil.

Ecophysiological responses of two poplar species to intraspecific and interspecific competition under different nitrogen levels

Aims Populus deltoides and P.euramericana are widely used in China as major forestry species.At present,little is known about their responses to nitrogen(N)deficiency when grown in monocultures or mixed plantations.The aim of this investigation was to analyze the growth,and morphological and physiological responses of P.deltoides and P.euramericana to different N levels under competition conditions.Methods We employed two Populus species(P.deltoides and P.euramericana)to discover how N deficiency affects plant traits under different competition types(P.deltoides×P.deltoides,intraspecific competition;P.euramericana×P.euramericana,intraspecific competition;P.deltoides×P.euramericana,interspecific competition).Potted seedlings were exposed to two N levels(normal N,N deficiency),and nitrogen-and competition-driven differences in growth,morphology and physiology were examined.Important Findings Under normal N conditions,interspecific competition significantly decreased the total root weight,root mass fraction(RMF),root–shoot ratio(R/S)and carbon/nitrogen ratio(C/N),and increased the leaf dry weight,leaf mass fraction and total leaf area of P.euramericana compared with intraspecific competition.The same conditions significantly affected the growth and morphological variables of P.deltoides,except for the dry weight of fine roots,R/S,specific leaf area,RMF,total nitrogen content and C/N compared with intraspecific competition.In addition,chlorophyll a(Chla),total chlorophyll(Tchl),carotenoid contents(Caro)and the carbon isotope composition(δ13C)of P.deltoides were significantly lower in interspecific competition than in intraspecific competition,but no difference was detected in P.euramericana.The effects of N deficiency on P.deltoides under intraspecific competition were stronger than under interspecific competition.In contrast,the effects of N deficiency on P.euramericana between intraspecific and interspecific competition were not significantly different.These results suggest that under normal N condition,P.deltoides is expected to gain an advantage in monocultures rather than in mixtures with P.euramericana.Under N deficiency,the growth performance of P.euramericana was more stable than that of P.deltoides under both cultivation modes.