Root responses to nitrogen pulse frequency under different nitrogen amounts

Responses of morphology and biomass allocation of roots to frequency of nitrogen (N) pulse potentially influence the fitness of plants, but such responses may be determined by root size. We grew 12 plant species of three functional groups (grasses, forbs, and legumes) under two N pulse frequencies (high vs. low supply frequency) and two N amounts (high vs. low supply amount). Compared to low-amount N supply, high-amount N supply stimulated biomass accumulation and root growth by either increasing the thickness and length of roots or decreasing the root mass fraction. Compared to low-frequency N supply, high-frequency N supply improved biomass accumulation and root growth in forbs or grasses, but not in legumes. Furthermore, the magnitude of the response to N frequency was significantly negatively correlated with root size at the species scale, but this was only true when the N amount was high. We conclude that root responses to N frequency are related to plant functional types, and non-legume species is more sensitive to N frequency than legume species. Our results also suggest that root size is a determinant of root responses to N frequency when N supply amount is high.     

Root morphological plasticity and nutrient acquisition of perennial grass species from habitats of different nutrient availability

We studied the root foraging ability and its consequences for the nutrient acquisition of five grass species that differ in relative growth rate and that occur in habitats that differ widely in nutrient availability. Foraging responses were quantified, based on the performance of the plants in homogeneous and heterogeneous soil environments of the same overall nutrient availability. Although all species tended to produce a significantly higher root length density in a nutrient-rich patch, this response was significant only for the faster-growing species. The increased root length density resulted from small, though not significant, changes in root biomass and specific root length. The effectiveness of root proliferation was determined by quantifying the total amount of nutrients (N and P) accumulated by the plants over the course of the experiment. Plants acquired more N in a heterogeneous environment than in a homogeneous environment, although the total nutrient availability was the same. The ability to acquire nutrients (N or P) in the heterogeneous environment was not related to the ability of species to increase root length density in response to local nutrient enrichment. In contrast to other studies, our results suggest that the role of morphological plasticity of roots in acquiring patchily distributed resources is limited. Possible reasons for this discrepancy are discussed. Received: 11 September 1997 / Accepted: 28 February 1998     

Growth and morphological responses to water level and nutrient supply in three emergent macrophyte species

Sanjiang Plain is the largest freshwater marsh in China, where plant zonation along water-level gradients is a common phenomenon. The aim of this experiment was to identify the role of water level and nutrient availability on plant zonation in the plain. Growth and root morphology of three perennial emergent macrophyte species were investigated by growing in two water levels (0.1 and 10.0 cm, relative to soil surface) and in two levels of nutrient supply (0 and 0.5 g slow-release fertilizer per container). In the plain, Carex lasiocarpa typically occurs at low elevations, Glyceria spiculosa at medial elevations, and Deyeuxia angustifolia at high elevations. The relative growth rate was the highest in C. lasiocarpa and the lowest in D. angustifolia in the 10.0-cm water level. Among the three species, only total biomass of D. angustifolia was affected by water level, and decreased with increasing water level. High nutrient supply led to increased total biomass in C. lasiocarpa and G. spiculosa. High water level led to an increased root diameter in G. spiculosa and a decreased root length in C. lasiocarpa. In the 10.0-cm water level, low nutrient supply led to thinner roots in D. angustifolia, but resulted in an increased specific root length (SRL) in C. lasiocarpa and root diameter in G. spiculosa. Water-level effect on root porosity was only observed in G. spiculosa, and nutrient amendment did not influence root porosity in all the species. These data indicate that both nutrient and water level are important factors regulating plant distribution pattern in the Sanjiang Plain, because both C. lasiocarpa and G. spiculosa are relatively sensitive to nutrient supply whereas D. angustifolia is sensitive to water level. Handling editor: S. M. Thomaz     

水曲柳和落叶松人工林根系分解与养分释放

采用埋袋法对水曲柳和落叶松粗根(5～10 mm)、中根(2～5 mm)、细根(＜2 mm)的分解速率及其养分释放进行了为期2年的研究.结果表明,水曲柳粗根、中根和细根年分解系数分别为0.3649、0.4381和0.2720,落叶松依次分别为0.1967、0.1955和0.2464.通过养分分析发现,根系分解过程释放大量C和养分.分解150 d后,两树种所有级别根系的可溶性糖释放均超过90%.水曲柳粗根和中根K的释放均在40%左右,细根K的释放为71%,落叶松所有级别的根系K的释放均在95%以上.在根系分解第2年,两树种粗根和中根N的释放在50%左右波动,P在40%左右波动,两树种细根N和P的释放均达到60%.因此,根系分解在C和养分循环中起重要作用,如果将其忽略,土壤有机质和养分元素的循环将会被严重低估.     

不同镉水平下小麦对镉及矿质养分吸收和积累的品种间差异

土壤Cd污染不仅影响作物的生长发育和产量形成,也对食物安全产生了潜在威胁,本试验以苗期生长特性有明显差异的2个小麦品种为供试材料,设置0、0．03、0．1、0．3和1．0mg.kg^-1Cd等5种处理水平,研究不同Cd水平下两品种的生长和Cd及矿质养分吸收,结果表明,低浓度Cd（0.03mg.kg^-1)对两品种的生长和干物质积累有明显的促进作用,而高浓度Cd（＞0.3mg.kg^-1)则显著抑制生长,且受抑制程度与品种有关,甘谷534的耐性相对较强;地上部和根系的Cd含量,Cd处理水平与品种之间存在着显著的互作,低Cd水平下（＜0.1mg.kg^-1),甘谷534的Cd含量较高,而高Cd水平下,则以鄂81513的Cd含量较高;Cd处理显著影响矿质养分的吸收,鄂81513在0.1mg.kg^-1Cd水平下5种大量元素含量明显低于对照,而甘谷534变化相对较小,这与两品种生长和分蘖对Cd处理的反应表现一致。     

Root growth and litter decomposition in a coffee plantation under shade trees

In a non-fertilized coffee plantation under shade trees the root biomass was excavated to estimate its distribution in the soil profile. One third of total fine (less than 1 mm) roots was found in the first 10 cm of soil; the cumulative total to 30 cm reached 73%. A highly variable and transient amount of fine roots colonized the litter layer. Root production both in the litter and in the first 7.5 cm of mineral soil was estimated from sequential samplings and was 10 g m^–2 yr^–1 and 660 g m^–2 yr^–1 respectively. The decomposition rate of weighed averages of litter fractions in the coffee plantation, calculated as the ratio of litter fall rate to the amount found in the soil was k=4.8. Shade tree leaves, the major component of litter descomposed slower than coffee leaves and these slower than flowers and fruits. Litter bag experiments showed considerable slower rates when mesh was 0.03 mm than 0.5 mm. Nitrogen and phosphorous showed increases in concentrations as decomposition progressed while potassium, calcium and magnesium followed a decrease in concentration that paralleled that of dry weight loss. In comparing the decomposition rate for litter with or without coffee roots growing in the bags, a tendency to show faster decomposition rates was found for the treatment with roots. These differences were however, only significant for one month for shade tree leaves litter. Nitrogen amounts remaining in shade tree leaves litter was lower in the treatment with roots that without roots. Potassium concentration in roots was positively correlated with potassium concentration in decomposing leaf litter where roots were growing. These results suggest that while roots growing attached to decomposing litter had little or no effect in speeding the decomposition process, the superficial roots seem to play an important role in absorbing very efficiently the mineralized nutrients from litter. The anatomical study of roots showed that the plantation is intensely infected with V-A mycorrhiza. External mycorrhizal hyphae did not to play a role in attachment of roots to decomposing litter while root hairs were found to grow in profusion on root surfaces oriented toward litter.     

Shifts in plant nutrient use strategies under secondary forest succession

In evergreen broad-leaved forests (EBLFs) in Tiantong National Forest Park, Eastern China, we studied the soil chemistry and plant leaf nutrient concentration along a chronosequence of secondary forest succession. Soil total N, P and leaf N, P concentration of the most abundant plant species increased with forest succession. We further examined leaf lifespan, leaf nutrient characteristics and root–shoot attributes of Pinus massoniana Lamb, the early-successional species, Schima superba Gardn. et Champ, the mid-successional species, and Castanopsis fargesii Franch, the late-successional species. These species showed both intraspecific and interspecific variability along succession. Leaf N concentration of the three dominant species increased while N resorption tended to decrease with succession; leaf P and P resorption didn’t show a consistent trend along forest succession. Compared with the other two species, C. fargesii had the shortest leaf lifespan, largest decay rate and the highest taproot diameter to shoot base diameter ratio while P. massoniana had the highest root–shoot biomass ratio and taproot length to shoot height ratio. Overall, P. massoniana used ‘conservative consumption’ nutrient use strategy in the infertile soil conditions while C. fargesii took up nutrients in the way of ‘resource spending’ when nutrient supply increased. The attributes of S. superba were intermediate between the other two species, which may contribute to its coexistence with other species in a wide range of soil conditions.     

Seed selection by dark-eyed juncos (Junco hyemalis): optimal foraging with nutrient constraints?

Summary Observations of the foraging behavior of six captive dark-eyed juncos (Junco hyemalis) are used to test the assumptions and predictions of optimal diet choice models (Pyke et al. 1977) that include nutrients (Pulliam 1975). The birds sequentially encountered single seeds of niger thistle (Guizotia abyssinica) and of canary grass (Phalaris canariensis) on an artificial substrate in the laboratory. Niger thistle seeds were preferred by all birds although their profitability in terms of energy intake (J/s) was less than the profitability of canary grass seeds. Of four nutritional components used to calculate profitabilities (mg/s) lipid content was the only characteristic that could explain the junco's seed preference. As predicted by optimal diet theory the probability of consuming niger thistle seeds was independent of seed abundance. However, the consumption of 71–84% rather than 100% of the seeds encountered is not consistent with the prediction of all-or-nothing selection. Canary grass seeds were consumed at a constant rate (no./s) independent of the number of seeds encountered. This consumption pattern invalidates a model that assumes strict maximization. However, it is consistent with the assumption that canary grass seeds contain a nutrient which is required in minimum amounts to meet physiological demands (Pulliam 1975). These experiments emphasize the importance of incorporating nutrients into optimal foraging models and of combining seed preference studies with studies of the metabolic requirements of consumers.     

Fine root growth and demographic responses to nutrient patches in four old-field plant species

Proliferation of roots in a nutrient patch can occur either as a result of an increase in root length (morphological response) or by a change in root birth or death rates (demographic responses). In this study we attempted to distinguish between these two mechanisms of response to nutrient patches and to compare the responses of four old-field plant species (two annuals, two perennials). For all four species combined, there were significant increases in root numbers and root length in fertilized patches. Root proliferation in fertilized patches was largely due to increased birth (=branching) rates of new roots. However, there was also a significant increase in root death rates in the fertilized patches which reduced the magnitude of the increase in net root numbers. Plots for individual species suggested they differed in the magnitude and timing of root proliferation in fertilized patches due to differences in root birth and death rates. However, because of the limited sample size in this study, there was only a marginally significant difference among species in root birth rates, and no difference in death rates. Further studies are currently underway to better quantify species differences in the demographic mechanism, as well as magnitude, of response to nutrient patches and if this would affect the ability to exploit small-scale heterogeneity in soil resources.     

Root Caps and Rhizosphere

In this paper we discuss recent work on the physiological, molecular, and mechanical mechanisms that underlie the capacity of root caps to modulate the properties of the rhizosphere and thereby foster plant growth and development. The root cap initially defines the rhizosphere by its direction of growth, which in turn occurs in response to gradients in soil conditions and gravity. The ability of the root cap to modulate its environment is largely a result of the release of exudates and border cells, and so provides a potential method to engineer the rhizosphere. Factors affecting the release of border cells from the outer surface of the root cap, and function of these cells and their exudates in the rhizosphere, are considered in detail. Release of border cells into the rhizosphere depends on soil matric potential and mechanical impedance, in addition to a host of other environmental conditions. There is good evidence of unidentified feedback signals between border cells and the root cap meristem, and some potential mechanisms are discussed. Root border cells play a significant mechanical role in decreasing frictional resistance to root penetration, and a conceptual model for this function is discussed. Root and border cell exudates influence specific interactions between plant hosts and soil organisms, including pathogenic fungi. The area of exudates and border cell function in soil is an exciting and developing one that awaits the production of appropriate mutant and transgenic lines for further study in the soil environment.     

Root growth and plant biomass in <Emphasis Type="Italic">Lolium perenne</Emphasis> exploring a nutrient-rich patch in soil

We investigated soil exploration by roots and plant growth in a heterogeneous environment to determine whether roots can selectively explore a nutrient-rich patch, and how nutrient heterogeneity affects biomass allocation and total biomass before a patch is reached. Lolium perenne L. plants were grown in a factorial experiment with combinations of fertilization (heterogeneous and homogeneous) and day of harvest (14, 28, 42, or 56 days after transplanting). The plant in the heterogeneous treatment was smaller in its mean total biomass, and allocated more biomass to roots. The distributions of root length and root biomass in the heterogeneous treatment did not favor the nutrient-rich patch, and did not correspond to the patchy distribution of inorganic nitrogen. Specific root length (length/biomass) was higher and root elongation was more extensive both laterally and vertically in the heterogeneous treatment. These characteristics may enable plants to acquire nutrients efficiently and increase the probability of encountering nutrient-rich patches in a heterogeneous soil. However, heterogeneity of soil nutrients would hold back plant growth before a patch was reached. Therefore, although no significant selective root placement in the nutrient-rich patch was observed, plant growth before reaching nutrient-rich patches differed between heterogeneous and homogeneous environments.     

The extracellular proteome of Bacillus licheniformis grown in different media and under different nutrient starvation conditions

The now finished genome sequence of Bacillus licheniformis DSM 13 allows the prediction of the genes involved in protein secretion into the extracellular environment as well as the prediction of the proteins which are translocated. From the sequence 296 proteins were predicted to contain an N-terminal signal peptide directing most of them to the Sec system, the main transport system in Gram-positive bacteria. Using 2-DE the extracellular proteome of B. licheniformis grown in different media was studied. From the approximately 200 spots visible on the gels, 89 were identified that either contain an N-terminal signal sequence or are known to be secreted by other mechanisms than the Sec pathway. The extracellular proteome of B. licheniformis includes proteins from different functional classes, like enzymes for the degradation of various macromolecules, proteins involved in cell wall turnover, flagellum- and phage-related proteins and some proteins of yet unknown function. Protein secretion is highest during stationary growth phase. Furthermore, cells grown in complex medium secrete considerably higher protein amounts than cells grown in minimal medium. Limitation of phosphate, carbon and nitrogen sources results in the secretion of specific proteins that may be involved in counteracting the starvation.     

匍匐茎草本活血丹(Glechoma longituba)在不同养分条件下的克隆形态

通过人工施肥的方法 ,对活血丹无性系实验种群在不同养分供应情况下的形态特征进行了研究。研究结果表明 :施肥可以显著改变克隆植株的形态 ,无性系分株产生能力增强 ,个体生长旺盛 ;匍匐茎总长增长 ,生物量增大 ,节间短 ,匍匐茎较粗壮 ;相对叶面积小 ,叶片厚 ,叶柄较粗短。在未施肥条件下 ,克隆植株形态表现则正好相反。养分供应的差异对叶片的形态以及根重和根量的影响较其对隔离者和叶柄的特征的影响更大。     

Root responses to legume plants integrate information on nitrogen availability and neighbour identity

Rhizobial symbiosis is known to increase the nitrogen availability in the rhizosphere of legumes. Therefore, it has been hypothesized that other plants’ roots should forage towards legume neighbours, but avoid non-legume neighbours. Yet, root distribution responding to legume plants as opposed to non-legumes has not yet been rigorously tested and might well be subject to integration of multiple environmental cues.In this study, wedevised an outdoor mesocosm experiment to examine root distributions of the two plant species Pilosella officinarum and Arenaria serpyllifolia in a two-factorial design. While one factor was ‘neighbour identity’, where plants were exposed to different legume or non-legume neighbours, the other factor was ‘nitrogen supply’. In the latter the nutrient-poor soil was supplemented with either nitrogen-free or with nitrogen-containing fertilizer.Unexpectedly, of all treatments that included a legume neighbour (eight different species or factor combinations), we found merely one case of root aggregation towards a legume neighbour (P. officinarum towards Medicago minima under nitrogen-fertilized conditions). In this very treatment, also P. officinarum root–shoot allocation was strongly increased, indicating that neighbour recognition is coupled with a contesting strategy.Considering the various response modes of the tested species towards the different legume and non-legume neighbours, we can conclude that roots integrate information on neighbour identity and resource availability in a complex manner. Especially the integration of neighbour identity in root decisions must be a vital aptitude for plants to cope with their complex biotic and abiotic environment in the field.     

匍匐茎草本活血丹(Glechomalongituba)在不同养分条件下的克隆形态

通过人工施肥的方法,对活血丹无性系实验种群在不同养分供应情况下的形态特征进行了研究。研究结果表明：施肥可以显著改变克隆植株的形态,无怀系分株产生能力增强,个体生长旺盛;匍匐茎总长增长,生物量增大,节间短,匍匐茎较粗壮;相对叶面积小,叶片厚,叶柄较粗短。在未施肥条件下,克隆植株形态表现则正好相反。养分供应的差异对叶片的形态以及根重和根量的影响较其对隔离者和叶柄的特征的影响更大。     

植物根毛的发生、发育及养分吸收

根毛是植物吸收养分的重要器官,认识根毛的发生、发育规律及其与养分吸收的关系,可为植物养分吸收效率的遗传改良提供依据.介绍了植物根毛的形态特性、发生和发育过程及其调控机制,并结合本实验室的工作,讨论了根毛对养分吸收的贡献、根毛受养分有效性的调节及其与其他根系形态构型性状间的关系,阐述了根毛中养分转运等植物营养过程及其生理和分子生物学基础.最后提出了关于根毛研究中的一些问题和研究前景.     

Induction of somatic embryogenesis and embryo development in Rumex acetosella L.

Axillary buds of the dioecious plant Rumex acetosella L. were isolated and cultured in vitro. The callus tissue which developed at the basal parts of the explants displayed a high capacity for shoot formation. This morphogenetic pattern was predominant on Murashige and Skoog (MS) medium supplemented with 2% sucrose, 2.2 mgl^-1 benzylaminopurine and 0.17 mgl^-1 indole-3-acetic acid. Somatic embryogenesis was induced when the osmolality of the medium was increased by adding 6% sucrose instead of 2%, or hexitols in addition to 2% sucrose. Most of the embryogenic calli were formed on the basal parts of leaf laminae and bracts. Development and maturation was strongly promoted by transferring the tissue to a solid or liquid medium lacking benzylaminopurine and indole-3-acetic acid and supplemented with 10 mgl^-1 gibberellic acid. The embryos germinated and developed into normal rosette plants when transferred to vermiculite moistened with hormone-free, half-strength MS salt solution. The histology of successive embryogenic stages is presented.     

Numerical study of some age-dependent parameters in root nutrient uptake

Computer simulation of root nutrient uptake has become a very powerfull tool in the analysis of plant and soil characteristics. One of the shortcomings of earlier numerical models is the lack of a proper accounting for age-dependent root parameters. In this article we present an algorithm that not only allows for time-and/or space-varying root growth rates, convective moisture uptake, root density, initial distribution of nutrient, effective diffusion coefficients, and buffer power; but also accounts for time-varying root efflux, root absorption power and maximum nutrient influxive rate.Several elementary numerical examples of NH ₄ ⁺ , NO ₃ ^– , P and K uptake for roots with temporally varying characteristics are presented.Contribution from the Purdue Agric. Exp. Stn., W. Lafayette, IN. Journal Paper No. 9476.     

Differences between resource patches modify root herbivore effects on plants

While soil resource heterogeneity and root herbivory can have significant direct influences on plant growth, soil heterogeneity may also have indirect effects by influencing the foraging behavior of root herbivores. We used sand-filled greenhouse pots to assess root herbivore foraging behavior and potential interactions between patch quality, herbivore foraging, and plant biomass production (yield). Individual pots were divided into four quarters: one fertilized, and three unfertilized, two of which were planted with tree seedlings. Two treatments were used to create fertilized quarters: high-organic manure fertilizer, and slow-release mineral fertilizer. Seedlings of red maple (Acer rubrum L.) and Virginia pine (Pinus virginiana L.) were used to create two single-species and one mixed-species treatments. Root-feeding beetle larvae were added to the pots and allowed to forage freely for ∼8 weeks. At harvest, root herbivores in organic-fertilized pots were strongly attracted to fertilized quarters despite their relatively low-root biomass. Herbivore distribution was significantly different in mineral fertilized pots, where larvae were most abundant in planted quarters, which is also where most of the plant roots occurred. Whole pot plant yield was significantly reduced by larvae; this effect was stronger in the mineral fertilized pots than organic fertilized pots. While one of the plant species appeared more sensitive to herbivory, root herbivores had a greater influence on yield in mixed-species pots than in single-species pots. Overall, these results suggest that patch quality influences on herbivore foraging may indirectly alter yield and plant community composition. Responsible Editor: Angela Hodge.     

Root biomass and nutrient dynamics in a scrub-oak ecosystem under the influence of elevated atmospheric CO2

Elevated CO₂ can increase fine root biomass but responses of fine roots to exposure to increased CO₂ over many years are infrequently reported. We investigated the effect of elevated CO₂ on root biomass and N and P pools of a scrub-oak ecosystem on Merritt Island in Florida, USA, after 7 years of CO₂ treatment. Roots were removed from 1-m deep soil cores in 10-cm increments, sorted into different categories (<0.25 mm, 0.25–1 mm, 1–2 mm, 2 mm to 1 cm, >1 cm, dead roots, and organic matter), weighed, and analyzed for N, P and C concentrations. With the exception of surface roots <0.25 mm diameter, there was no effect of elevated CO₂ on root biomass. There was little effect on C, N, or P concentration or content with the exception of dead roots, and <0.25 mm and 1–2 mm diameter live roots at the surface. Thus, fine root mass and element content appear to be relatively insensitive to elevated CO₂. In the top 10 cm of soil, biomass of roots with a diameter of <0.25 mm was depressed by elevated CO₂. Elevated CO₂ tended to decrease the mass and N content of dead roots compared to ambient CO₂. A decreased N concentration of roots <0.25 mm and 1–2 mm in diameter under elevated CO₂ may indicate reduced N supply in the elevated CO₂ treatment. Our study indicated that elevated CO₂ does not increase fine root biomass or the pool of C in fine roots. In fact, elevated CO₂ tends to reduce biomass and C content of the most responsive root fraction (<0.25 mm roots), a finding that may have more general implications for understanding C input into the soil at higher atmospheric CO₂ concentrations.