向日葵种群中植株个体大小对其氮素利用策略的影响

我们利用Berendse和Aerts提出的氮素利用效率(NUE)概念及原理研究了高密度一年生草本植物向日葵(Helianthus annuus L.)种群中植株个体大小对其氮素吸收利用的影响,并对种内竞争进行了分析。结果表明,植株对氮素的吸收与其个体大小不成线性关系,说明种群内不同植株个体对土壤氮素的竞争属于非对称竞争。植株的氮素损失随着个体大小的增加而增加。个体较大的植株具有较高的氮素输入率和较低的氮素输出率,因而具有较高的氮素净增加值。植株的氮素生产力(NP)和氮素平均滞留时间(MRT)均与植株个体大小呈正相关。较大的植物个体具有较高的NP和较长的MRT,由于NUE为NP和MRT二者的乘积,因而较大个体植株的NUE高于个体较小的植株。同种植物的不同个体的NP和MRT之间不存在协衡关系。氮素回收效率(NRE)与植株个体大小密切相关。在个体水平上,较大的植株个体具有较高的NUE与其较高的NRE有关。种群内植株个体对土壤氮素的非对称竞争主要由于植株对氮素的吸收和利用效率不同所致。因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分利用策略,对于种内不同植株的养分策略研究也同样适用。     

藜个体在高密度种群中的氮素利用效率

 氮素利用效率（NUE）是植物养分策略研究中的一项重要内容。该文利用Berendse和Aerts提出的氮素利用效率概念和原理研究了高密度的藜(Chenopodium album)种群中不同植物个体在种内竞争条件下的氮素利用效率。结果表明,由于植株的氮素吸收速率与其个体大小成非线性关系,说明不同植株个体对氮素的竞争属于非对称竞争。个体较大的植株氮素输入较高,而个体较小的植株氮素输出较高,因而较大个体植株的氮素净增加也较高。植株的氮素损失随着个体大小的增加而增加,较大植株个体的氮素浓度随着生长而下降,而较小植株个体的氮素浓度随时间的变化不大,说明个体较小的植株的生长受光照的限制比受氮素的限制更大,而对较大的植株个体而言,它们的生长受氮素的限制更大。高密度藜种群中的不同植物个体具有不同的养分策略,氮素利用效率及其组成部分氮素生产力（NP）和氮素滞留时间（MRT）均不同。植株的NP和MRT与其个体大小正相关,较大的植物个体具有较高的NP和较长的MRT,因而氮素利用效率也高于个体较小的植株。在个体水平上,种内不同植株的NP与MRT不存在权衡关系（Trade-off）。因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分策略,对于研究种内不同植株的养分策略也同样适用。     

盐胁迫下施肥对棉花生长及氮素利用的影响

利用海水配制不同含盐量(0、0.15%、0.3%)的土壤盆栽棉花,在可移动遮雨棚内研究了不同施肥(N、NK、NP、NPK)处理对棉花生长、氮素吸收与利用的影响.结果表明: 盐胁迫和施肥均影响棉花生物产量、棉株氮素农学利用效率、氮素生物利用效率和氮素积累量,且两者存在显著的互作效应.施肥能提高盐胁迫下棉株氮素利用效率及氮素积累量,并显著增产,不同施肥处理中以N、P、K肥料配合施用的效果最好;施肥效果受盐胁迫程度的影响,低盐胁迫(0.15%)下的施肥效果好于中度盐胁迫(0.3%).     

C3和C4植物的氮素利用机制

提高植物的氮素利用效率(NUE)不仅有利于保障全球粮食安全, 也是实现农业可持续发展的重要途径。近半个世纪以来, 植物氮素利用机理研究已取得重要进展, 但NUE的调控机制仍不明确, NUE的提高仍然十分有限。高等植物集光合碳素同化和氮素同化于一体, 只有碳氮代谢相互协调, 才能维持植物体内的碳氮平衡, 保证植物正常生长发育。由于C₃和C₄植物的光合氮素利用率(PNUE)存在差异, 对氮素的利用效率也会存在差异。为了更有效地提高作物的NUE, 须更全面地了解C₃和C₄植物对氮素吸收、转运、同化和信号转导等关键因子的功能和调控机制。此外, 面对大气CO₂浓度增高和全球气候变暖条件下的植物碳氮同化及其机理的研究也不容忽视。该文综述了C₃和C₄植物氮素利用关键因素的差异及其调控机制, 并对提高C₃禾本科作物氮素利用效率的遗传改良途径进行了展望。     

南方水稻氮素吸收与利用效率的基因型差异及评价

 以南方籼型水稻（Oryza sativa）品种为试验材料进行大田试验,以探讨提高水稻氮素吸收与利用效率的基因型潜力。结果表明,除早季分蘖期氮素积累量、干物质生产效率和抽穗期氮素积累量以及晚季氮素运转效率外,各基因型氮素吸收与利用效率存在显著或极显著的差异,提高水稻氮素吸收与利用效率的基因型潜力很大。基因型生育期对其氮素吸收与利用效率产生重要影响,生育期较长的基因型其氮素吸收效率、稻谷和干物质生产效率以及农艺效率较高。杂交稻氮素的生产效率、农艺效率、回收效率和收获指数较常规稻高,但二系杂交稻并没有比三系杂交稻明显提高。通过排序方式对各基因型氮素吸收与利用效率进行评价的结果表明,不同氮素吸收与利用效率指标的排序以及同一指标早晚季的排序均存在较大差异。氮素吸收与利用效率经标准化后的综合排序可对各基因型的氮素吸收与利用效率进行综合评价,吻合系数则可较好地反映各基因型早晚季氮素吸收与利用效率的排序。     

施氮量对小麦氮磷钾养分吸收利用和产量的影响

高产条件下研究了不同施氮量对小麦植株氮、磷、钾养分吸收利用及籽粒产量的影响.结果表明,适量施氮可促进小麦植株对氮素的吸收与积累,较高的施氮量不利于起身期之后的氮素积累,致使成熟期小麦氮素积累量未能显著提高;与不施氮肥相比,施氮显著提高植株磷素积累量;随施氮量增加,植株磷素积累量增加不显著;施氮量增加促进小麦生育前期对钾素的吸收积累,在生育后期降低植株钾素的流失.随施氮量增加,籽粒氮素含量呈先增后降的趋势,氮素向籽粒的分配比例趋于降低,植株氮素利用效率无显著变化,氮素收获指数下降;不同施氮处理之间籽粒磷素含量和钾素含量无显著差异,施氮量增加,营养器官钾素含量、钾素积累量和钾素向叶片的分配比例均呈增加趋势;同时,磷素和钾素利用效率降低;不同施氮处理间,植株磷素、钾素收获指数无显著差异.籽粒产量随施氮量增加呈先增加后降低的趋势,以施氮195 kg/hm2的处理籽粒产量最高.     

中国东部南北样带森林优势植物叶片的水分利用效率和氮素利用效率

通过测定中国东部南北样带主要森林生态系统中10种优势植物(兴安落叶松、蒙古栎、水曲柳、紫椴、色木槭、红松、杉木、木荷、马尾松、锥栗)叶片的碳氮含量(Cmass、Nmass)、同位素丰度(δ13C、δ15N)以及光合响应曲线,分析了不同优势植物叶片的水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明： 不同生活型植物叶片的Nmass和δ15N差异显著,表现为阔叶植物>针叶植物,落叶植物>常绿植物;最大光合速率(Pn max)表现为针叶植物>阔叶植物,落叶植物>常绿植物;植物叶片的瞬时水分利用效率(WUEi)和长期水分利用效率(WUE)均表现为阔叶植物>针叶植物,常绿植物>落叶植物;植物叶片的瞬时氮素利用效率(NUEi)和长期氮素利用效率(NUE)则表现出相反的规律,且常绿植物和落叶植物叶片的NUE差异显著;WUEi和WUE之间相关性不显著,而NUEi和NUE之间呈显著正相关.植物叶片的水分利用效率与氮素利用效率显著负相关.两种资源利用效率均受植物生活型的影响,并且存在一定的制约关系.     

不同水氮水平下小麦品种对光、水和氮利用效率的权衡

通过再裂区设计田间试验,以3个春小麦品种(和尚头、西旱2号和宁春4号)为材料,设置两个灌溉水平(充分灌水4500 m³·hm^-2和有限灌水3000 m³·hm^-2)和5个施氮水平(0、75、150、225、300 kg N·hm^-2),研究小麦光能利用效率(LUE)、水分利用效率(WUE)、氮素利用效率(NUE)对水氮的响应特性及其相互关系.结果表明: 3个小麦品种间LUE、WUE和NUE差异显著.在一定范围内增加灌水和施氮量则LUE升高,过量施氮则LUE下降.强抗旱和中等抗旱品种(和尚头和西旱2号)WUE受灌水量的影响比不抗旱品种(宁春4号)小.施氮可以调节小麦WUE,中等施氮水平(和尚头和西旱2号在150 kg N·hm^-2时,宁春4号在225 kg N·hm^-2时)有最高的WUE.随施氮量增加,植株氮素累积量先增后减,氮素干物质生产效率(NUE_b)、氮素收获指数(NHI)、氮肥农学利用效率(NAE)和氮肥偏生产力(PFP)均显著降低.灌溉水平对NHI无显著影响;随灌水量增加,小麦氮素积累量显著增加,强抗旱和中等抗旱品种NUE_b和NAE显著降低,不抗旱品种 NUE_b和PFP显著升高,对其他指标无显著影响.3个小麦品种氮素获取能力与氮素利用效率呈极显著负相关,NUE_b与LUE、WUE呈显著负相关,LUE与WUE呈显著正相关,春小麦氮素利用效率与光能利用效率、水分利用效率间存在明显的权衡关系.当灌水量为3000 m³·hm^-2,强抗旱和中等抗旱品种在150 kg N·hm^-2,不抗旱品种在225 kg N·hm^-2时,有较高的资源利用效率.     

添加玉米残体对土壤-植物系统中氮素转化的影响

采用盆栽试验和^15N示踪技术对黑土添加玉米残体(秸秆和根茬)土壤-植物系统中氮素转化进行了研究,结果表明,玉米残体还田能够增加土壤氮素含量,减轻因其作为燃烧材料而造成的氮素损失和对大气的污染,玉米残体施入土壤,增加了土壤微生物氮含量,提高土壤氮活性,有利于土壤氮素养分的协调供应,玉米残体配施氮肥与氮肥单施相比,玉米植株氮素累积量相近,但氮素在玉米植株不同器官中的分配比例不同;添加玉米残体能够促进氮素从营养器官向籽粒中转移,提高氮素养分的利用效率,同时,添加玉米残体还可以降低土壤NO^-3-N的累积,减少肥料氮的损失4．7％～5．6％。     

植物养分高效利用机制研究进展

长期进化和环境适应导致不同植物或同种植物不同基因型间养分利用效率(NUE)差异明显,研究筛选植物养分高效利用基因型至关重要,其极大的增产潜力可补充代替传统植物栽培方法所需的能源.目前人们对于植物NUE概念的理解存在一定差异,造成众多研究成果缺乏可比性.通过对植物NUE的概念及其描述方式难以统一原因的分析,提出人工林NUE应采用干材生物量与林分养分总量的比值表示.综合评述了植物体内养分高效利用及植物对生长介质中养分高效吸收的生物生理学适应性机制.对养分逆境植物养分高效利用适应性策略的整个过程进行了描述,进一步阐明了Ca2+在化学通讯机制中的生物功能,指出Ca2+可能是启动植物养分高效利用挽救机制的主要调控因子,并就该领域今后研究工作的特点作了展望.     

Nitrogen uptake and use by competing individuals in a Xanthium canadense stand

We studied differences in nitrogen uptake and use for plant growth among individuals competing in a natural dense stand of an annual herb, Xanthium canadense. Larger individuals took up more nitrogen than proportionately to their size, indicating that the competition for soil nitrogen was asymmetric among individuals, although it was more symmetric than the competition for light. The rate of nitrogen loss of individuals also increased with plant size. While smaller individuals shared smaller fractions of total plant nitrogen in the stand, they had higher nitrogen concentrations per unit mass. "Turnover" rates of nitrogen influx (r_in) and outflux (r_out) were defined as the rates of nitrogen uptake and loss per unit aboveground nitrogen, respectively. r_in was higher in larger individuals, whereas r_out was higher in smaller individuals. Consequently, the relative rate of nitrogen increment (r_in-r_out) was higher in larger individuals, whereas it was around zero in the smallest individuals. The mean residence time of nitrogen (MRT), defined as the inverse of r_out, was longer in larger individuals. Nitrogen productivity (NP), i.e. the growth rate per unit aboveground nitrogen, was higher in larger individuals. As the product of lifetime MRT and NP gives the nitrogen use efficiency (NUE), defined as biomass production per unit flux of nitrogen, higher MRT and NP observed in larger individuals would have contributed to their higher lifetime NUE. Shorter MRT in smaller individuals was caused by the abscission of leaves which contained relatively large fractions of total plant nitrogen. Xanthium canadense, as a competitive ruderal, tended to produce leaves at higher positions to acquire higher light levels at the expense of older leaves rather than to modify their productive structure to efficiently use low light levels as observed in shade-tolerant species.     

Effect of Nitrogen Supply on the Nitrogen Use Efficiency of an Annual Herb, Helianthus annuus L.

Nitrogen use efficiency (NUE) is the product of nitrogen productivity (NP) and the mean residence time of nitrogen (MRT). Theory suggests that there should be a trade-off between both components,but direct experimental evidence is still scarce. To test this hypothesis, we analyzed the effect of varying nitrogen supply levels on NUEand its two components (NP, MRT) in Helianthus annuus L., an annual herb.The plants investigated were subjected to six nitrogen levels (0, 2, 4, 8, 16, and 32 g N/m2). Total plant production increased substantially with increasing nitrogen supply. Nitrogen uptake and loss also in creased with nitrogen supply. Nitrogen influx (rin) and outflux (rout) were defined as the rates of nitrogen uptake and loss per unit aboveground nitrogen, respectively. Both rin and rout increased with increasing nitrogen supply. In addition, rin was far higher than rout. Consequently, the relative rate of nitrogen incre ment (rin- rout) also increased with nitrogen supply. There were marked differences between treatments with respect to parameters related to the stress resistance syndrome: nitrogen pool size, leaf nitrogen concentration,and net aboveground productivity increased with nitrogen supply. Plants at high nitrogen levels showed a higher NP (the growth rate per unit aboveground nitrogen) and a shorter MRT (the inverse of rout), whereas plants at low nitrogen levels displayed the reverse pattern. Shorter MRT for plants at high nitrogen levels was caused by the abscission of leaves that contained relatively large fractions of total plant nitrogen. We found a negative relationship between NP and MRT, the components of NUE, along the gradient of nitrogen availability, suggesting that there was a trade-off between NP and MRT. The NUE increased with increasing nitrogen availability, up to a certain level, and then decreased. These results offer support for the hypoth esis that adaptation to infertile habitats involves a low nitrogen loss (long MRT in the plant) rather than a high NUE per se. The higher NUE at the plant level was a result, in part, of greater nitrogen resorption during senescence. We suggest that a long MRT (an index of nitrogen conservation) is a potentially successful strategy in nitrogen-poor environments.     

Leaf-level nitrogen use efficiency: definition and importance

Nitrogen use efficiency (NUE) has been widely used to study the relationship between nitrogen uptake and dry mass production in the plant. As a subsystem of plant nitrogen use efficiency (NUE), I have defined leaf-level NUE as the surplus production (gross production minus leaf respiration) per unit amount of nitrogen allocated to the leaf, with factorization into leaf nitrogen productivity (NP) and mean residence time of leaf nitrogen (MRT). These concepts were applied to two herbaceous stands: a perennial Solidago altissima stand and an annual Amaranthus patulus stand. S. altissima had more than three times higher leaf NUE than A. patulus due to nearly three times longer MRT of leaf N. In both species, NUE and NP were higher at the leaf level than at the plant level, because most leaf N is involved directly in the photosynthetic activity and because leaf surplus production is higher than the plant net production. MRT was longer at the plant level. The more than twice as long MRT at the plant level as at the leaf level in S. altissima was due to a large contribution of nitrogen storage belowground in the winter in this species. Thus, comparisons between a perennial and an annual system and between plant- and leaf-level NUE with their components revealed the importance of N allocation, storage, recycling, and turnover of organs for leaf photosynthetic production and plant dry mass growth.     

Nitrogen use efficiency revisited

Nitrogen use efficiency (NUE) was originally defined as the dry mass productivity per unit N taken up from soil. The term was subsequently redefined as the product of nitrogen productivity (NP) and mean residence time of nitrogen (MRT). However, this redefinition was found to contradict the original definition under certain conditions, and confusion arose when the MRT defined for a steady-state system was applied to a system that was actually not at steady state. As MRT is the expected length of time that a unit of N newly taken up from soil is retained before being lost, it can be translated into the plant nitrogen duration (PND) divided by the total N uptake. This MRT is determined equally well for a steady state- and a non-steady state system and is in accordance with the original definition of NUE. It can be applied to a herbaceous perennial stand (that was at a steady state) and to an annual stand (that was not at a steady state) to determine NUE. NUE is also applicable when plant growth and reproduction are analyzed in relation to N use.     

Growth and nitrogen use in Xanthium canadense grown in an open or in a dense stand

Plants develop branches profusely when grown solitarily, while less so when grown in a dense stand. Such changes in architecture are associated with changes in dry mass allocation and nitrogen use. Here, we studied what traits in plant growth and nitrogen use were influenced by different light climates in the stand. Annual plants (Xanthium canadense) were grown solitarily or in a dense stand. Dry mass growth was analyzed as the product of the net assimilation rate (NAR) and leaf area (LA). Nitrogen use efficiency (NUE) was analyzed as the product of nitrogen productivity (NP) and the mean residence time (MRT) of nitrogen. These growth variables were further factorized into their components. Solitary plants maintained a high NAR, whereas plants in the dense stand decreased the NAR due to mutual shading. Plants in the dense stand developed a larger LA with a higher specific leaf area than solitary plants. Solitary plants had higher NUE due to higher NP. A temporal increase in NUE was attributed to the increase in MRT of nitrogen. Light climate was different between solitary and dense-stand plants, but they took up a comparable amount of nitrogen and used it differently in response to the given light climate. NUE was thus demonstrated to be a useful tool for analyzing the mechanism leading to different N use in plant growth.     

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.     

Nitrogen-use efficiency in six perennial grasses from contrasting habitats

1. We studied the nitrogen-use efficiency (NUE) in six perennial grasses adapted to a wide range of nutrient availability. The glasshouse experiment was carried out in pots containing nutrient solution, with two fertility treatments. Nitrogen-use efficiency was considered as the product of nitrogen productivity and mean residence time of the nitrogen in the plant (calculated using ¹⁵N pulse labelling).
2. The species investigated are characteristic of habitats ranging from very nutrient rich to extremely nutrient poor, in the following order: Lolium perenne, Arrhenatherum elatius, Festuca rubra, Anthoxanthum odoratum, Festuca ovina and Molinia caerulea .
3. Lolium perenne (adapted to nutrient-rich habitats) had higher nitrogen productivity ( A ) than M. caerulea (species adapted to nutrient-poor habitats) but lower than that of F. rubra (from habitats with an intermediate availability of nutrients).
4. In the low fertility treatment, species with the lowest nitrogen-use efficiency had the lowest N productivity and the highest mean nitrogen residence time (MRT); however, although species with the highest nitrogen use efficiency had the highest N productivity they did not have the lowest MRT. In all species the nitrogen-use efficiency decreased with increasing N supply. The two components of the NUE ( A and MRT) are inversely correlated along gradients of nutrient availability, but not at very high levels of nutrient availability.
5. The nitrogen-use efficiency of species at constant levels of nutrient supply tends to increase with increasing nutrient availability in their preferred habitat, according to the Clausman nutrient index, up to a certain nutrient availability and then decreases. The results support the contention that species from nutrient-poor sites are not necessarily adapted by a high nitrogen-use efficiency, but by low nutrient loss rates (high mean residence time of N in the plant).     

Does leaf shedding increase the whole-plant carbon gain despite some nitrogen being lost with shedding?

When old leaves are shed, part of the nitrogen in the leaf is retranslocated to new leaves. This retranslocation will increase the whole-plant carbon gain when daily C gain : leaf N ratio (daily photosynthetic N-use efficiency, NUE) in the old leaf, expressed as a fraction of NUE in the new leaf, becomes lower than the fraction of leaf N that is resorbed before shedding (R(N)). We examined whether plants shed their leaves to increase the whole-plant C gain in accord with this criterion in a dense stand of an annual herb, Xanthium canadense, grown under high (HN) and low (LN) nitrogen availability. The NUE of a leaf at shedding expressed as a fraction of NUE in a new leaf was nearly equal to the R(N) in the LN stand, but significantly lower than the R(N) in the HN stand. Thus shedding of old leaves occurred as expected in the LN stand, whereas in the HN stand, shedding occurred later than expected. Sensitivity analyses showed that the decline in NUE of a leaf resulted primarily from a reduction in irradiance in the HN stand. On the other hand, it resulted from a reduction in irradiance and also in light-saturated photosynthesis : leaf N content ratio (potential photosynthetic NUE) in the LN stand.