Mapping QTLs for root traits under different nitrate levels at the seedling stage in maize (Zea mays L.)

Nitrogen (N) loss is a worldwide problem in crop production. Apart from reasonable N fertilizer application, breeding N efficient cultivars provides an alternative way. Root architecture is an important factor determining N acquisition. However, little is known about the molecular genetic basis for root growth in relation to N supply. In the present study, an F₈ maize (Zea may L.) recombinant inbred (RI) population consisting of 94 lines was used to identify the QTLs for root traits under different nitrate levels. The lateral root length (LRL), axial root length (ARL), maximal axial root length (MARL), axial root number (ARN) and average axial root length (AARL) were evaluated under low N (LN) and high N (HN) conditions in a hydroponics system. A total of 17 QTLs were detected among which 14 loci are located on the same chromosome region as published QTLs for root traits. A major QTL on chromosome 1 (between bnlg1025 and umc2029) for the AARL under LN could explain 43.7% of the phenotypic variation. This QTL co-localizes with previously reported QTLs that associate with root traits, grain yield, and N uptake. Our results indicate that longer axial roots are important for efficient N acquisition and the major QTL for AARL may be used as a marker in breeding N efficient maize genotypes.     

玉米幼苗地上部/根间氮的循环及其基因型差异

以两个玉米（ZeamaysL.）自交系原引1号（YY1）和综31（Z31）为研究材料,采用盆栽土培的培养方法,在正常供氮（HN,0.15gN/kg干土）和低氮量供应（LN,0.038gN/kg干土）培养条件下对玉米幼苗植株体内氮的循环量及其在地上部/根间的分配量进行了定量地测定、计算。结果表明,在玉米幼苗地上部/根间氮的循环量很高。低氮量供应使玉米幼苗植株吸氮量下降,根中氮的分配比例增加,同时地上部/根间氮的循环量也随之减少。与氮低效自交系Z31相比,氮高效自交系YY1幼苗中地上部/根间的氮循环量大、氮向根的分配量高,因而有利于其根系的生长,表现为根/地上部之比和总根长较高。这可能有利于其中后期对氮素的高效吸收与利用。     

Impacts of maize hybrids with different nitrogen use efficiency on root-associated microbiota based on distinct rhizosphere soil metabolites

Plant genotypes shape root-associated microbiota that affect plant nutrient acquisition and productivity. It is unclear how maize hybrids modify root-associated microbiota and their functions and relationship with nitrogen use efficiency (NUE) by regulating rhizosphere soil metabolites. Here, two N-efficient (NE) (ZD958, DMY3) and two N-inefficient (NIE) maize hybrids (YD9953, LY99) were used to investigate this issue under low N (60 kg N ha⁻¹, LN) and high N (180 kg N ha⁻¹, HN) field conditions. NE hybrids had higher yield than NIE hybrids under LN but not HN. NE and NIE hybrids recruited only distinct root-associated bacterial microbiota in LN. The bacterial network stability was stronger in NE than NIE hybrids. Compared with NIE hybrids, NE hybrids recruited more bacterial taxa that have been described as plant growth-promoting rhizobacteria (PGPR), and less related to denitrification and N competition; this resulted in low N₂O emission and high rhizosphere NO₃⁻-N accumulation. NE and NIE hybrids had distinct rhizosphere soil metabolite patterns, and their specific metabolites were closely related to microbiota and specific genera under LN. Our findings reveal the relationships among plant NUE, rhizosphere soil metabolites, root-associated microbiota, and soil nutrient cycling, and this information is informative for breeding NE crops.     

A novel morphological response of maize (Zea mays) adult roots to heterogeneous nitrate supply revealed by a split‐root experiment

Approximately 35–55% of total nitrogen (N) in maize plants is taken up by the root at the reproductive stage. Little is known about how the root of an adult plant responds to heterogeneous nutrient supply. In this study, root morphological and physiological adaptations to nitrate‐rich and nitrate‐poor patches and corresponding gene expression of ZmNrt2.1 and ZmNrt2.2 of maize seedlings and adult plants were characterized. Local high nitrate (LoHN) supply increased both lateral root length (LRL) and density of the treated nodal roots of adult maize plants, but only increased LRL of the treated primary roots of seedlings. LoHN also increased plant total N acquisition but not N influx rate of the treated roots, when expressed as per unit of root length. Furthermore, LoHN markedly increased specific root length (m g^?1) of the treated roots but significantly inhibited the growth of the lateral roots outside of the nitrate‐rich patches, suggesting a systemic carbon saving strategy within a whole root system. Surprisingly, local low nitrate (LoLN) supply stimulated nodal root growth of adult plants although LoLN inhibited growth of primary roots of seedlings. LoLN inhibited the N influx rate of the treated roots and did not change plant total N content. The gene expression of ZmNrt2.1 and ZmNrt2.2 of the treated roots of seedlings and adult plants was inhibited by LoHN but enhanced by LoLN. In conclusion, maize adult roots responded to nitrate‐rich and nitrate‐poor patches by adaptive morphological alterations and displayed carbon saving strategies in response to heterogeneous nitrate supply.     

Effects of nutrient availability on water hyacinth standing crop and detritus deposition

Nutrient-enriched water hyacinths were stocked in outdoor tanks and cultured under both high nutrient (HN) and low nutrient (LN) regimes for 10 months. Seasonal changes in standing crop biomass and morphology of LN water hyacinths were similar to those of HN water hyacinths, despite a ten-fold between-treatment difference in N availability and a two-fold difference in average plant N concentrations (1.0 and 2.0% for LN and HN plants, respectively). Tissue N accumulated by the LN plants prior to stocking helped support standing crop development during the 10 month study. In both HN and LN treatments, the rate of detritus deposition, or the sloughing of dead plant tissues from the mat, was lower than the actual detritus production rate because of the retention of dead ‘aerial’ tissues (laminae and petioles) in the floating mat. The retention of laminae and petioles may serve as a nutrient conservation mechanism, since nutrients released from decomposing tissues in the mat-water environment may be assimilated by adjacent plants. The average rate of detritus deposition (both dry matter and N) by LN water hyacinths (1.2 g dry wt. m⁻² day⁻¹ and 0.017 g N m⁻² day⁻¹) was lower than that of HN plants (3.0 g dry wt. m⁻² day⁻¹ and 0.075 g N m⁻² day⁻¹) during the study. Low detrital N losses by the water hyacinth probably enhance the survival of this species in aquatic systems which receive nutrient inputs intermittently.     

Changes in root size and distribution in relation to nitrogen accumulation during maize breeding in China

Background and aims

Modern maize breeding has increased maize yields worldwide. The changes in above-ground traits accompanying yield improvement are well-known, but less information is available as to the effect of modern plant breeding on changes in maize root traits.

Methods

Root growth, nitrogen uptake, dry matter accumulation and yield formation of six maize hybrids released from 1973 to 2000 in China were compared. Experiments were conducted under low and high nitrogen supply in a black soil in Northeast China in 2010 and 2011.

Results

While nitrogen accumulation, dry matter production and yield formation have been increased, modern maize breeding in China since 1990 has reduced root length density in the topsoil without much effect on root growth in the deeper soil. The efficiency of roots in acquiring N has increased so as to match the requirement of N accumulation for plant growth and yield formation. The responses of root growth, nitrogen and dry matter accumulation, and grain yield to low-N stress were similar in the more modern hybrids as in the older ones.

Conclusions

Modern maize breeding has constitutively changed root and shoot growth and plant productivity without producing any specific enhancement in root responsiveness to soil N availability.     

模拟氮沉降对天山云杉细根分解及其养分释放的影响

采用野外模拟试验,设计4种氮处理——对照(不施氮,CK)、低氮(施氮5kg·hm-2·a-1,LN)、中氮(施氮10kg·hm-2·a-1,MN)、高氮(施氮15kg·hm-2·a-1,HN),研究氮沉降对天山云杉细根分解及养分释放的影响。结果表明:(1)不同氮处理分解2年后天山云杉细根残留率依次为74.044%(HN)、71.967%(MN)、68.156%(CK)、61.933%(LN),且差异显著。(2)天山云杉的细根月分解速率在试验前期不同氮处理下规律不明显;而在试验后期呈现为对照中氮低氮高氮。(3)4种氮处理下天山云杉细根分解50%需要的时间依次为3.31年(LN)、3.67年(CK)、4.28年(MN)、4.64年(HN),分解95%需要的时间依次为14.39年(LN)、15.93年(CK)、18.58年(MN)和20.17年(HN)。(4)天山云杉细根C元素迁移模式总体表现为直接释放,N元氮为富集-释放模式,残留率呈现波动式下降趋势。(5)不同氮处理下天山云杉细根分解率与C元素浓度间均呈线性负相关关系;对照和低氮处理下,天山云杉细根分解率与N元素浓度间均为线性负相关关系,中氮和高氮处理下,细根分解率随N元素浓度的增加呈先增加后降低的趋势。     

东北地区4种林分土壤呼吸及温、湿度敏感性对氮添加的短期响应

全球变化中氮沉降日益严重,已对森林生态系统的各个过程产生了重要影响。因此,通过研究氮添加对森林生态系统土壤碳输出的影响,对分析全球变化背景下土壤碳吸存具有重要意义。对黑龙江省帽儿山实验林场白桦(Betula platyphylla)次生林,以及水曲柳(Fraxinus mandschurica)、红松(Pinus koraiensis)、长白落叶松(Larix olgensis)人工林通过2年氮添加(对照(0 kg N hm~(-2) a~(-1)),低氮(50 kg N hm~(-2) a~(-1)),中氮(100 kg N hm~(-2) a~(-1))和高氮(150 kg N hm~(-2) a~(-1)))试验,测定根生物量密度、土壤微生物量碳浓度、土壤呼吸速率及温、湿度敏感性等指标,旨在探讨森林生态系统土壤呼吸对氮添加的短期响应。结果表明:(1)低氮处理对白桦和水曲柳林土壤呼吸速率影响不显著,但显著提高了红松和长白落叶松林土壤呼吸速率;水曲柳林分中高氮处理土壤呼吸速率显著降低于低氮和中氮处理,而其他林分高氮处理土壤呼吸速率仅显著低于低氮处理。(2)氮添加处理下,4种林分中林分土壤呼吸速率与根生物量密度呈极显著正相关,Pearson相关系数为0.81。(3)低氮处理下5 cm和10 cm处土壤呼吸温度敏感性系数Q_(10)值较CK处理分别提高了2.65%和3.12%,高氮处理较CK处理分别降低了6.29%和5.46%。但氮添加处理对土壤呼吸和土壤湿度间的相关性无影响。综上所述,阔叶林与针叶林土壤呼吸速率对氮添加的响应存在差异。根生物量密度是影响不同林分土壤呼吸对短期氮添加响应的主要因素,同时氮添加处理显著改变了土壤温度敏感性系数。     

Effects of sugars and growth regulators on <Emphasis Type="Italic">in vitro</Emphasis> growth of <Emphasis Type="Italic">Dactylorhiza</Emphasis> species

The influence of sugars and growth regulators on shoot and root growth of Dactylorhiza species was studied under in vitro conditions. The seedling development was stimulated with the application of glucose and sucrose at concentration of 10 g dm⁻³ each. The improvement of shoot growth rate and shoot length was enhanced by cytokinins N ⁶-(2-isopentenyl)adenine or N ⁶-benzyladenine and their combination with auxin indolebutyric acid (IBA). The root growth rate and root length of seedlings increased in the presence of IBA and α-naphthaleneacetic acid. Individual Dactylorhiza species showed statistically significant differences in shoot and root development depending on sugar and growth regulator combinations.     

Effects of enhanced atmospheric ammonia on photosynthetic characteristics of two maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.) cultivars with various nitrogen supply across long-term growth period and their diurnal change patterns

We investigated the effect of enhanced atmospheric ammonia (NH₃) in combination with low and high nitrogen (LN and HN, respectively) growth medium on photosynthetic characteristics of two maize (Zea mays L.) cultivars (NE5 with high- and SD19 with low N-use efficiency) across long-term growth period and their diurnal change patterns exposed to 10 nl l⁻¹ and 1,000 nl l⁻¹ NH₃ fumigation in open-top chambers (OTCs). Regardless of the level of N in medium, increased NH₃ concentration promoted maximum net photosynthetic rate (P _max) and apparent quantum yield (AQY) of both cultivars at earlier growth stages, but inhibited P _max of NE5 from silking to maturity stage and that of SD19 at maturity stage only above the ambient concentration. Greater positive/less negative responses were predominant in the LN than in the HN treatment, especially for SD19. Dark respiration rate (R _D) remained more enhanced in the LN than in the HN treatment for SD19 as well as increased in the LN while decreased in the HN treatment for NE5 at their silking stage, following exposure to elevated NH₃ concentration. Additionally, enhanced atmospheric NH₃ increased net photosynthetic rate (P _N) and stomatal conductance (g _s) but reduced intercellular CO₂ concentration (C _i) of both cultivars with either the LN or HN treatment during the diurnal period at tasseling stage. The diurnal change patterns of P _N and g _s showed bimodal curve type and those of C _i presented single W-curve type for NE5, when NH₃ concentration was enhanced. As for SD19, single-peak curve type was showed for both P _N and g _s while single V-curve type for C _i. All results supported the hypothesis that appropriately enhanced atmospheric NH₃ can increase assimilation of CO₂ by improving photosynthesis of maize plant, especially at earlier growth stages and after photosynthetic “noon-break” point. These impacts of elevated NH₃ concentration were more beneficial for SD19 as compared to those for NE5, especially in the LN supply environment.     

Morphological compatibility of white clover and perennial ryegrass cultivars grown under two nitrate levels in flowing solution culture

The effects of nitrate (NO3-) supply on shoot morphology, vertical distribution of shoot and root biomass and total nitrogen (N) acquisition by two perennial ryegrass (Lolium perenne L.) cultivars (AberElan and Preference) and two white clover (Trifolium repens L.) cultivars (Grasslands Huia and AberHerald) were studied in flowing nutrient culture. Cultivars were grown from seed as monocultures and the clovers inoculated with Rhizobium. The 6-week measurement period began on day 34 (grasses) and day 56 (clovers) when the NO3- supply was adjusted to either 2 mmol m-3 (low nitrogen, LN) or 50 mmol m-3 (high nitrogen, HN). These treatments were subsequently maintained automatically. Plants were harvested at intervals to measure their morphology and N content. Cultivars of both species differed significantly in several aspects of their response to NO3- supply. In the grasses, the LN treatment increased the root : shoot ratio of AberElan but did not affect the distribution of root length in the root profile. In contrast, this treatment changed the root distribution of Preference compared with HN, resulting in a larger proportion of root length being distributed further down the root profile. The morphology of white clover Grasslands Huia was for the most part unaffected by the level of NO3- supply. In contrast, AberHerald exhibited different growth strategies, with LN plants increasing their stolon weight per unit length at the expense of leaf production, leaf area and stolon length, whereas HN plants showed reduced stolon thickness, greater leaf area production and stolon length per plant. Cultivars with different morphological/physiological strategies in response to NO3- supply may be of value in the construction of 'compatible mixtures' aimed at reducing oscillations in sward clover content by extending the range of conditions that allow balanced coexistence of species to occur.     

A comprehensive analysis of root morphological changes and nitrogen allocation in maize in response to low nitrogen stress

The plasticity of root architecture is crucial for plants to acclimate to unfavourable environments including low nitrogen (LN) stress. How maize roots coordinate the growth of axile roots and lateral roots (LRs), as well as longitudinal and radial cell behaviours in response to LN stress, remains unclear. Maize plants were cultivated hydroponically under control (4 mm nitrate) and LN (40 μm ) conditions. Temporal and spatial samples were taken to analyse changes in the morphology, anatomical structure and carbon/nitrogen (C/N) ratio in the axile root and LRs. LN stress increased axile root elongation, reduced the number of crown roots and decreased LR density and length. LN stress extended cell elongation zones and increased the mature cell length in the roots. LN stress reduced the cell diameter and total area of vessels and increased the amount of aerenchyma, but the number of cell layers in the crown root cortex was unchanged. The C/N ratio was higher in the axile roots than in the LRs. Maize roots acclimate to LN stress by optimizing the anatomical structure and N allocation. As a result, axile root elongation is favoured to efficiently find available N in the soil.     

Seedling responses to band applied NH4OH rates and to N form in two maize hybrids

Growth of maize seedlings can be improved by enhanced ammonium nutrition, but placing fertilizer anhydrous ammonia close to seedlings introduces the risk of ammonia toxicity. In this study, growth and root elongation response to rates of closely placed NH₄OH bands were investigated in two contrasting maize hybrids. Seven rates of NH₄OH, ranging from 0 to 200 mg N kg^-1 soil were injected into the center of each pot. A single rate of Ca(NO₃)₂-N was included to compare hybrids for N form preference at a moderate N rate. Three seedlings per pot were planted 5.7 cm from the injection point.Hybrid B73×LH51 produced a quadratic response in shoot growth to NH₄OH rates, whereas LH74×LH123 exhibited a significant linear decline in response to NH₄OH rate. Root length density sampled from the fertlized zone declined linearly in response to NH₄OH rate while a slight increase in root length density in unfertilized zones was observed at intermediate NH₄OH rates. Hybrids did not differ in root length density in either zone.The hybrid with greater tolerance of NH₄OH rates (B73×LH51) also showed a preference in shoot growth for NH₄-over NO₃-N at 66.7 mg N kg^-1 compared to LH74×LH123. On average across hybrids, nitrate concentrations of xylem exudate collected from detopped plants were 14.5 mmol g^-1 for Ca(NO₃)₂ treatments and 1.5 mmol g^-1 for NH₄OH treatments, indicating that contrasting N-form nutrition resulted from fertilizer treatments. Malate concentrations were higher in the NH₄OH treatment indicating that this organic acid anion may substitute for the negative charge of nitrate during enhanced ammonium nutrition in maize.The results suggest that potentially useful genetic variation exists in maize for N form preference and for tolerance to increasing ammonical-N rates.     

Carbon and nitrogen partitioning during the post‐anthesis period is conditioned by N fertilisation and sink strength in three cereals

Further knowledge of the processes conditioning nitrogen use efficiency (NUE) is of great relevance to crop productivity. The aim of this paper was characterise C and N partitioning during grain filling and their implications for NUE. Cereals such as bread wheat (Triticum aestivum L. cv Califa sur), triticale (× Triticosecale Wittmack cv. Imperioso) and tritordeum (× Tritordeum Asch. & Graebn line HT 621) were grown under low (LN, 5 mm NH₄NO₃) and high (HN, 15 mm NH₄NO₃) N conditions. We conducted simultaneous double labelling (¹²CO₂ and ¹⁵NH₄¹⁵NO₃) in order to characterise C and N partitioning during grain filling. Although triticale plants showed the largest total and ear dry matter values in HN conditions, the large investment in shoot and root biomass negatively affected ear NUE. Tritordeum was the only genotype that increased NUE in both N treatments (NUE_total), whereas in wheat, no significant effect was detected. N labelling revealed that N fertilisation during post‐anthesis was more relevant for wheat and tritordeum grain filling than for triticale. The study also revealed that the investments of C and N in flag leaves and shoots, together with the ‘waste’ of photoassimilates in respiration, conditioned the NUE of plants, and especially under LN. These results suggest that C and N use by these plants needs to be improved in order to increase ear C and N sinks, especially under LN. It is also remarkable that even though tritordeum shows the largest increase in NUE, the low yield of this cereal limits its agronomic value.     

Mapping QTLs for nitrogen uptake in relation to the early growth of wheat (Triticum aestivum L.)

The objective of this study was to map QTLs for N uptake (NUP) in wheat, and to investigate factors influencing NUP. Two independent field trials with low N (LN) and high N (HN) treatments were conducted in the growing seasons of 2002–2003 (trial 1) and 2003–2004 (trial 2) to measure NUP per plant (N accumulated in the aerial part at maturity stage) of a doubled haploid (DH) population consisting of 120 DH lines derived from winter wheat varieties Hanxuan 10 and Lumai 14. A hydroponic culture with all nutrients supplied sufficiently was conducted to investigate shoot dry weight (SDW), root dry weight (RDW), tiller number (TN) and NUP (total plant N uptake) per plant of this mapping population at seedling stage. SDW, RDW, TN and NUP investigated in the hydroponic culture were significantly and positively correlated with each other, and with NUP under both LN and HN conditions in the field trials. Nine and eight QTLs for NUP were detected under LN and HN conditions in the field trials, respectively. Four to five QTLs for SDW, RDW, TN and NUP were detected in the hydroponic culture. One SDW QTL, three RDW QTLs, two TN QTLs detected in the hydroponic culture were linked with QTLs for NUP under LN or HN condition in the field trials. The positive correlation and genetic linkage for the traits between the field trials and the hydroponic culture demonstrated that greater seedling vigor of root and shoot is an important factor influencing N uptake in wheat. Diaoguo An and Junying Su: These authors contributed equally to this work. Section Editor: H.J. Kronzucker     

模拟氮沉降对杉木幼苗细根的生理生态影响

细根对氮沉降的生理生态响应将显著影响森林生态系统的生产力和碳吸存。为了揭示氮沉降对杉木细根的生理生态影响,对一年生杉木(Cunninghamia lanceolata)幼苗进行了模拟氮沉降试验,并测定施氮1年后杉木幼苗细根生物量、细根形态学特征(比根长、比表面积)、元素化学计量学指标(C、N、P、C/N、C/P、N/P)、细根代谢特征(细根比呼吸速率、非结构性碳水化合物)。结果表明:(1)杉木细根生物量随氮添加水平的升高而显著降低,尤其是0—1 mm细根生物量;细根比根长和比表面积随氮添加水平升高而显著增大。(2)氮添加后杉木细根C含量、C/N、C/P显著降低,高氮添加导致1—2 mm细根N含量和N/P显著升高,而低氮添加导致1—2 mm细根P含量显著升高、N/P显著降低,而0—1 mm细根的N、P含量则保持相对稳定。(3)氮添加后杉木细根比呼吸速率无显著变化,细根可溶性糖含量随氮添加增加而显著增加,而淀粉含量和NSC显著降低。综合以上结果表明:氮添加后用于细根形态构建的碳分配减少,这可能会减少土壤中有机碳的保留,0—1 mm细根的形态更易发生变化,但是其内部N、P养分含量相对更稳定以维持生理活动,细根NSC对氮添加的响应表明施氮可能导致细根受光合产物的限制。     

Growth and allocation by a keystone wetland plant, <Emphasis Type="Italic">Panicum hemitomon</Emphasis>, and implications for managing and rehabilitating coastal freshwater marshes,Louisiana, USA

This paper attempts to establish linkages between growth by a keystone wetland plant, Panicum hemitomon Schultes, and the independent and interactive effect of nutrient and hydrologic regime to inform management and rehabilitation of thick-mat floating marsh (TMFM). To do so a manipulative glasshouse experiment employing created TMFM similar to that under consideration for field trials and two levels each of N, P and hydrology was conducted. P. hemitomon grew vigorously under saturated (flooding level with the surface of the mat) when compared to inundated (+15 cm flooding) hydrologic conditions, and under enriched (50 g m⁻² year⁻¹) when compared to non-enriched (25 g m⁻² year⁻¹) N. Further, and as inferred from net CO₂ assimilation, shoot biomass and rhizome biomass and length, N-enriched conditions seemed to lessen inundation stress. For all variables the interaction between N and hydrology was non-significant and there was no observable effect of P. We were unable to infer root or mat buoyancy from root specific gravity measurements but it was evident at harvest that saturation or minimal flooding is required for vigorous root and rhizome growth. This study provides insight to the notion that decreased mat buoyancy (and increased flooding level) resulting from sediment deposition associated with Mississippi River diversions could adversely affect TMFM sustainability, but more clearly demonstrates the need to maintain saturated hydrologic conditions for achieving the type of root and rhizome growth we feel is required for TMFM rehabilitation.     

Nitrogen remobilization response to current supply in young citrus trees

Internal nitrogen (N) storage and remobilization processes support seasonal growth (flowering/fructification and subsequent leaf development) in particular in early spring, when soil temperatures are unfavourable for adequate N uptake. Storage nitrogen mobilization in young citrus trees was studied under two contrasting N supplies; high N (HN) and low N dose (LN) in the critical period of flowering and fruit set. ¹⁵N labelling technique was used to distinguish N derived from internal remobilization from that taken up by the roots. Regardless N supply, the greatest N remobilization took place from the beginning of the vegetative activity until flowering. Low N availability significantly increased (+14%) N retranslocation at the end of June drop agreeing with the hypothesis that reserve mobilization depends on soil N availability during flowering and fruit set. At the end of fruit drop, N remobilization contributed up to 70% and 61% of total N of young organs for LN and HN, respectively. Remobilized N was mainly recovered in abscised organs of both HN and LN trees and to a lesser extent in new flush leaves; however a greater percentage partitioned to abscised organs of LN as a consequence of the greater remobilization rate and the increased fruit abscission. Old leaves of LN remobilized significantly higher N, while woody organs and root system did not show differences between HN and LN supplied trees. The results presented in this paper demonstrate that the amount of N remobilized by young citrus plants depends on external N availability. Thus, low N application rates in early stages (flowering and fruit set) lead to higher translocation of N stored during the previous cycle to developing new organs.     

Competition for nitrogen between Pinus sylvestris and ectomycorrhizal fungi generates potential for negative feedback under elevated CO2

We investigated fungal species-specific responses of ectomycorrhizal (ECM) Scots pine (Pinus sylvestris) seedlings on growth and nutrient acquisition together with mycelial development under ambient and elevated CO₂. Each seedling was associated with one of the following ECM species: Hebeloma cylindrosporum, Laccaria bicolor, Suillus bovinus, S. luteus, Piloderma croceum, Paxillus involutus, Boletus badius, or non-mycorrhizal, under ambient, and elevated CO₂ (350 or 700 μl l⁻¹ CO₂); each treatment contained six replicates. The trial lasted 156 days. During the final 28 days, the seedlings were labeled with ¹⁴CO₂. We measured hyphal length, plant biomass, ¹⁴C allocation, and plant nitrogen and phosphorus concentration. Almost all parameters were significantly affected by fungal species and/or CO₂. There were very few significant interactions. Elevated CO₂ decreased shoot-to-root ratio, most strongly so in species with the largest extraradical mycelium. Under elevated CO₂, ECM root growth increased significantly more than hyphal growth. Extraradical hyphal length was significantly negatively correlated with shoot biomass, shoot N content, and total plant N uptake. Root dry weight was significantly negatively correlated with root N and P concentration. Fungal sink strength for N strongly affected plant growth through N immobilization. Mycorrhizal fungal-induced progressive nitrogen limitation (PNL) has the potential to generate negative feedback with plant growth under elevated CO₂. Responsible Editor: Herbert Johannes Kronzucker     

Seasonal changes in the effects of free-air CO2 enrichment (FACE) on growth, morphology and physiology of rice root at three levels of nitrogen fertilization

Over time, the relative effects of elevated [CO₂] on the aboveground photosynthesis, growth and development of rice (Oryza sativa L.) are likely to be changed with increasing duration of CO₂ exposure, but the resultant effects on rice belowground responses remain to be evaluated. To investigate the impacts of elevated [CO₂] on seasonal changes in root growth, morphology and physiology of rice, a free‐air CO₂ enrichment (FACE) experiment was performed at Wuxi, Jiangsu, China, in 2002–2003. A japonica cultivar with large panicle was exposed to two [CO₂] (ambient [CO₂], 370 μmol mol⁻¹; elevated [CO₂], 570 μmol mol⁻¹) at three levels of nitrogen (N): low (LN, 15 g N m⁻²), medium (MN, 25 g N m⁻²) and high N (HN, 35 g N m⁻²). Elevated [CO₂] increased cumulative root volume, root dry weight, adventitious root length and adventitious root number at all developmental stages by 25–71%, which was mainly associated with increased root growth rate during early growth period (EGP) and lower rate of root senescence during late growth period (LGP), while a slight inhibition of root growth rate occurred during middle growth period (MGP). For individual adventitious roots, elevated [CO₂] increased average length, volume, diameter and dry weight early in the season, but the effects gradually disappeared in subsequent stages. Total surface area and active adsorption area per unit root dry weight reached their maxima 10 days earlier in FACE vs. ambient plants, but both of them together with root oxidation ability per unit root dry weight declined with elevated [CO₂] during MGP and LGP, the decline being larger during MGP than LGP. The CO₂‐induced decreases in specific root activities during MGP and LGP were associated with a larger amount of root accumulation during EGP and lower N concentration and higher C/N ratio in roots during MGP and LGP in FACE vs. ambient plants. The results suggest that most of the CO₂‐induced increases in shoot growth of rice are similarly associated with increased root growth.