Limited costs of wrong root placement in Rumex palustris in heterogeneous soils

Nutrient hot spots in the soil have a limited life span, but the costs and benefits for root foraging are still underexposed. We assessed short-term costs that may arise when a nutrient-rich patch induces root proliferation, but then rapidly disappears. Rumex palustris plants were grown with a homogeneous or a heterogeneous nutrient application. After root proliferation in a nutrient-rich patch, nutrient supply was switched from homogeneous to heterogeneous, and vice versa, or the patch location was changed. R. palustris proliferated its roots in the rich patch. After switching, the relative growth rates of the roots were adjusted to the novel pattern of nutrient availability. However, the changes in local root biomass lagged behind the rapid shift in nutrient supply, because the root mass realized in specific sectors could not be rapidly relocated. Despite this, R. palustris did not exhibit costs of switching in terms of biomass or nitrogen uptake. Our data suggest that rapid shifts in uptake rate and redistribution of nitrogen within the plant may have lowered the costs of incorrect root placement.     

Dynamic phenotypic plasticity for root growth in Polygonum: a comparative study

Species differences in patterns of phenotypic plasticity may be an important aspect of adaptive diversity. Plasticity for functionally important root traits was studied in inbred field lineages of Polygonum persicaria and P. cespitosum (Polygonaceae). Replicate seedlings were grown in plexiglass rhizotrons under a range of constant and temporally variable moisture treatments. Plasticity was determined for final whole-plant biomass, root biomass allocation, and absolute and proportional root length. The dynamic aspect of root plasticity was examined by digitizing weekly tracings of the proportional deployment of each plant's root system to different vertical soil layers. Plants of both species expressed significant functionally adaptive phenotypic plasticity in the relative allocation, length, and vertical deployment of root systems in response to contrasting moisture conditions. Plasticity patterns in these closely related species were in general qualitatively similar, but for most traits differed in the magnitude and/or the timing of the plastic response. Dynamic changes in root deployment were more marked as well as faster in P. persicaria. Species differences in patterns of individual plasticity were generally consistent with the broader ecological distribution of P. persicaria in diverse as well as temporally variable moisture habitats.     

Signalling mechanisms underlying the morphological responses of the root system to nitrogen in Arabidopsis thaliana

Plants display considerable developmental plasticity in response to changing environmental conditions. The adaptations of the root system to variations in N supply are an excellent example of such developmental plasticity. In Arabidopsis, four morphological adaptations to the N supply have been characterized: (i) a localized stimulatory effect of external nitrate on lateral root elongation; (ii) a systemic inhibitory effect of high tissue nitrate concentrations on the activation of lateral root meristems; (iii) a suppression of lateral root initiation by high C:N ratios, and (iv) an inhibition of primary root growth and stimulation of root branching by external L-glutamate. These responses have provided valuable experimental systems for the study of N signalling in plants. This article will highlight some recent progress made in this direction from studies using the Arabidopsis root system. One recent development of note has been the emerging evidence of a regulatory role of nitrate transporters in some of the responses. It has been reported that the AtNRT1.1 (CHL1) dual-affinity nitrate transporter acts upstream of the ANR1 MADS box gene in mediating the stimulatory effect of a localized nitrate supply on lateral root proliferation. The AtNRT2.1 high-affinity nitrate transporter seems to be involved in the repression of lateral root initiation by high C:N ratios. The systemic inhibitory effect of high nitrate supply on lateral root development, which is mediated by abscisic acid (ABA), may be linked to the recently identified ABA receptor, FCA. The newly discovered root architectural response to external L-glutamate potentially offers a valuable experimental tool for studying the biological function of plant glutamate receptors and amino acid signalling.     

植物根系养分捕获塑性与根竞争

为了更有效地从土壤中获取养分, 植物根系在长期的进化与适应中产生了一系列塑性反应, 以响应自然界中广泛存在的时空异质性。同时, 植物根系的养分吸收也要面对来自种内和种间的竞争。多种因素都会影响植物根竞争的结果, 包括养分条件、养分异质性的程度、根系塑性的表达等。竞争会改变植物根系的塑性反应, 比如影响植物根系的空间分布; 植物根系塑性程度差异也会影响竞争。已有研究发现根系具有高形态塑性和高生理塑性的植物在长期竞争过程中会占据优势。由于不同物种根系塑性的差异, 固定的对待竞争的反应模式在植物根系中可能并不存在, 其响应随竞争物种以及土壤环境因素的变化而变化。此外, 随着时间变化, 根系塑性的反应及其重要性也会随之改变。植物对竞争的反应可能与竞争个体之间的亲缘关系有关, 有研究表明亲缘关系近的植物可能倾向于减小彼此之间的竞争。根竞争对植物的生存非常重要, 但目前还没有研究综合考虑植物的各种塑性在根竞争中的作用。另外根竞争对群落结构的影响尚待深入的研究。     

Physiological and morphological responses of red alder and sitka alder to flooding

Red alder (Alnus rubra Bong.) and sitka alder (A. viridis ssp. sinuata [Regel] Löve & Löve) are nitrogen-fixing woody species that grow sympatrically along the Pacific coast of North America. Red alder is found in poorly drained lowlands, as well as in soils of moist upland slopes, whereas sitka alder generally colonizes well-drained soils. To identify factors that contribute to flood tolerance, we conducted greenhouse experiments subjecting both species to a 20-day flood and 10-day recovery and red alder to a 50-day flood and 20-day recovery. We determined the effect of this stress on nitrogenase activity, root and nodule alcohol dehydrogenase (ADH) activity, lenticel and adventitious root development, relative growth rate (RGR), and leaf gas exchange. After 24 h of flooding, nitrogenase activity could not be detected in either species. Limited nitrogenase activity did return in red alder at the end of a 10-day recovery following the 20-day flood, but sitka alder showed no recovery of nitrogenase activity. After 50 days of continuous flooding, red alder nitrogenase activity returned to pretreatment levels. Red alder root and nodule ADH activity was more than twice that of sitka alder under flooded conditions. Sitka alder showed extensive root mortality and leaf abscission over the same 20-day flooding period. Flooded red alder exhibited an initial decline in root RGR, but recovered between days 10 and 20 with the formation of adventitious roots. Furthermore, initiation of adventitious roots in red alder coincided with an increase in stomatal conductance without a similar recovery of carbon dioxide exchange rate. Sitka alder formed few adventitious roots, lost much of its root and leaf biomass, and showed no restoration of growth during flooding or recovery. Different responses of red and sitka alder to flooding serve as a partial explanation for the different patterns of distribution of these species and suggest some adaptations of red alder that permit flood tolerance.     

Nitrogen uptake and root morphological responses of defoliated Lolium perenne (L.) to a heterogeneous nitrogen supply

Lolium perenne L. (c.v. Magella) plants were grown under three nutrient treatments for six weeks and then defoliated to test the hypothesis that for their regrowth they could acquire N equally well irrespective of N distribution. Two different N levels were applied; uniform level 1 N (U1), uniform level 2 (U2) and heterogeneous level 2 (H2). A system where the nutrient patch could be applied without barriers to root growth was adopted. A single defoliation to 4 cm height resulted in a reduction in tillering, biomass increment and N uptake at 3 weeks after defoliation. Root growth was reduced by defoliation under all N treatments. Defoliation was found to reduce the proportion of N in the shoots which was derived from root uptake from 7 to 14 days. At 21 days this effect was significant for the plants with a heterogeneously distributed supply only. By the end of the regrowth period, the undefoliated plants from H2 had a shoot biomass and N content equal to that of plants receiving the same total N but supplied homogeneously (U2). However, defoliation reduced the ability of the plant to acquire N from the patch. No preferential root growth was measured into the N-rich patch, but an increased root diameter within the patch was found. Root diameter was reduced by defoliation, coinciding with a reduction in concentration of N in the root tissue. As a result of the increased sink strength of the growing leaves after defoliation, the roots may become a source of carbon and also nitrogen. These responses to an N-rich patch under defoliation could alter a plant's competitive balance in a mixed sward. This revised version was published online in June 2006 with corrections to the Cover Date.     

不同养分环境下木荷种源生长和根系发育对邻株竞争响应的差异

研究同质和异质养分环境中邻株竞争对3个木荷种源生长和根系发育的影响,揭示了不同种源木荷生长竞争能力差异原因.结果表明: 与同质养分环境相比,3个木荷种源在异质养分环境中具有苗高生长量大、干物质积累量高和根系增生明显等特点.在异质养分环境下,福建建瓯种源木荷苗木生长量显著高于浙江龙泉和江西信丰种源,与杉木混植时尤为突出,这与其根系形态可塑性高和拓殖能力强有关,混植时,福建建瓯种源木荷的根长、根表面积和根体积等根系生长量较单植显著增长20.4 %～69.0%,其根系在富养表层大量增生的同时快速向深层贫养层拓殖,占有了更多的空间和资源,提高了觅养能力,使其生长优势更为明显,而浙江龙泉和江西信丰种源根系生长和深层土壤根系分布受邻株竞争影响不同程度的降低.纯植时,可能由于根系自我识别作用,3个木荷种源的根系生长发育均受到抑制,导致福建建瓯种源苗木生长量显著减小,而浙江龙泉种源苗高和干物质积累增长明显,这与其根系生理可塑性有关.建议生产上选用觅养效率高和竞争能力强的福建建瓯种源木荷,采用混交造林的方式提高木荷人工林生产力.     

Fine root responses to temporal nutrient heterogeneity and competition in seedlings of two tree species with different rooting strategies

There is little direct evidence for effects of soil heterogeneity and root plasticity on the competitive interactions among plants. In this study, we experimentally examined the impacts of temporal nutrient heterogeneity on root growth and interactions between two plant species with very different rooting strategies: Liquidambar styraciflua (sweet gum), which shows high root plasticity in response to soil nutrient heterogeneity, and Pinus taeda (loblolly pine), a species with less plastic roots. Seedlings of the two species were grown in sandboxes in inter‐ and intraspecific combinations. Nutrients were applied in a patch either in a stable (slow‐release) or in a variable (pulse) manner. Plant aboveground biomass, fine root mass, root allocation between nutrient patch and outside the patch, and root vertical distribution were measured. L. styraciflua grew more aboveground (40% and 27% in stable and variable nutrient treatment, respectively) and fine roots (41% and 8% in stable and variable nutrient treatment, respectively) when competing with P. taeda than when competing with a conspecific individual, but the growth of P. taeda was not changed by competition from L. styraciflua. Temporal variation in patch nutrient level had little effect on the species’ competitive interactions. The more flexible L. styraciflua changed its vertical distribution of fine roots in response to competition from P. taeda, growing more roots in deeper soil layers compared to its roots in conspecific competition, leading to niche differentiation between the species, while the fine root distribution of P. taeda remained unchanged across all treatments. Synthesis. L. styraciflua showed greater flexibility in root growth by changing its root vertical distribution and occupying space of not occupied by P. taeda. This flexibility gave L. styraciflua an advantage in interspecific competition.     

Growth and root responses of woody species to rocky substrate: implications for gully rehabilitation

Gullies formed in the Velhas River basin in Brazil have been filled with urban construction waste for physical stabilisation purposes. Aimed at rehabilitating gullies, we selected woody species from the Brazilian Cerrado that can grow on rocky substrates under greenhouse conditions. An assessment was made regarding plant growth in both rocky and natural soil substrates by analysing the height, diameter, fresh and dry weights of shoots and roots, plant water content, root occupation and architecture. Principal component analysis and Chi-squared tests segregated rock-tolerant species based on the specific influence on root dry and fresh weights. Fast-growing species reduced the emergence of their lateral roots under rocks, compromising their growth in height and biomass production. In contrast, slow-growing woody species were particularly suitable for gully rehabilitation because these species exhibited a genetic pattern of low lateral root emergence that prevented damage to their roots. Most slow-growing species demonstrated a similar growth pattern in both substrates, and some of them, such as Copaifera langsdorffii, achieved better growth in height and biomass production on rocks than on soil, a finding attributed to the root plastic response involving primary root elongation and lateral root emergence. Therefore, slow-growing species are recommended for gully rehabilitation procedures.     

Signaling mechanisms integrating root and shoot responses to changes in the nitrogen supply

During their life cycle, plants must be able to adapt to wide variations in the supply of soil nitrogen (N). Changes in N availability, and in the relative concentrations of NO₃ ⁻and NH₄ ⁺, are known to have profound regulatory effects on the N uptake systems in the root, on C and N metabolism throughout the plant, and on root and shoot morphology. Optimising the plant’s responses to fluctuations in the N supply requires co-ordination of the pathways of C and N assimilation, as well as establishment of the appropriate allocation of resources between root and shoot growth. Achieving this integration of responses at the whole plant level implies long-distance signaling mechanisms that can communicate information about the current availability of N from root-to-shoot, and information about the C/N status of the shoot in the reverse direction. In this review we will discuss recent advances which have contributed to our understanding of these long-range signaling pathways.     

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     

The effect of flooding and flood timing on leaf litter breakdown rates and nutrient dynamics in a river red gum (Eucalyptus camaldulensis) forest

Comparisons of litter standing-stocks in low-lying and higher areas of the floodplain and the effects of controlled flooding events on leaf litter decomposition and leaf litter nutrients were examined during autumn and winter in a southeastern Australian river red gum (Eucalyptus camaldulensis) floodplain forest. The mean mass of total litter and some litter components was significantly greater in autumn than in winter but there were few differences in litter mass between low-lying flood runners and higher sites (1.5 m) on the floodplain, regardless of season. Leaf decomposition was more rapid in flooded areas than in non-flooded areas and was significantly faster in autumn than in winter. In flooded leaves, concentrations of phosphorus and nitrogen dropped rapidly during the first 3 days of each experiment, increased to near original after 7–10 weeks and then decreased again. After 112 days of decomposition the C:N:P ratios of leaf litter increased, but this effect was most marked for flooded leaves. Simple models of leaf litter dynamics indicated that leaf litter standing-stocks in low-lying flood runners would be reduced by flooding, particularly during autumn. In contrast, models predicted a net gain in standing-stocks of leaf litter to be higher on the floodplain, particularly in autumn. Alteration to the seasonal timing of floods by river regulation has probably decreased litter standing-stocks and nutrients available in low-lying areas of the floodplain to support the production of macrophytes and biofilms during winter and spring floods.     

Influence of root herbivory on plant communities in heterogeneous nutrient environments

While plant species respond differently to nutrient patches, the forces that drive this variability have not been extensively examined. In particular, the role of herbivory in modifying plant-resource interactions has been largely overlooked. We conducted a glasshouse study in which nutrient heterogeneity and root herbivory were manipulated, and used differences in foraging among plant species to predict the influence of root herbivores on these species in competition. We also tracked the influence of neighborhood composition, heterogeneity, and herbivory on whole-pot plant biomass. When herbivores were added to mixed-species neighborhoods, Eupatorium compositifolium, the most precise forager, was the only plant species to display a reduction in shoot biomass. Neighborhood composition had the greatest influence on whole-pot biomass, followed by nutrient heterogeneity; root herbivory had the smallest influence. These results suggest that root herbivory is a potential cost of morphological foraging in roots. Root herbivores reduced standing biomass and influenced the relative growth of species in mixed communities, but their effect was not strong enough at the density examined to overwhelm the bottom-up effects of resource distribution.     

植物根系对氮胁迫的形态学响应

NO3-不仅是植物营养的主要N源,而且也是调节植物新陈代谢和生长发育的信号。植物根系对N供给的形态学适应表现在4个方面:(1)植物侧根(LR)响应体外硝酸盐的局部刺激而伸长,AtNRT1.1在侧根响应体外硝酸盐的局部刺激中,作用于ANR1 MADS box基因上游;(2)植物LR分裂组织活动受到组织中高浓度的硝酸盐抑制,RNA结合蛋白FCA是高硝酸盐/ABA诱导侧根发育抑制的信号传输途径成分;(3)植物侧根的发生受到体外高C∶N比抑制,AtNRT2.1参与高C∶N比抑制侧根发生;(4)植物根响应体外L-谷氨酸盐的刺激而分枝,初生根的生长受到体外L-谷氨酸盐抑制,植物根系对L-谷氨酸盐的响应可能与一种同源于哺乳动物离子型谷氨酸盐受体的植物蛋白的感知作用有关。植物根形态对N供给的响应具有重要生理和生态意义。     

Differences in the metabolic responses of root tips of wheat and rye to aluminium stress

The effects of aluminium (Al) ions on the metabolism of root apical meristems were examined in 4-day-old seedlings of two cereals which differed in their tolerance to Al: wheat cv. Grana (Al-sensitive) and rye cv. Dakowskie Nowe (Al tolerant). During a 24 h incubation period in nutrient solutions containing 0.15 mM and 1.0 mM of Al for wheat and rye, respectively, the activity of first two enzymes in the pentose phosphate pathway (G-6-PDH and 6-PGDH) decreased in the sensitive cultivar. In the tolerant cultivar activities of these enzymes increased initially, then decreased slightly, and were at control levels after 24 h. In the Al-sensitive wheat cultivar a 50% reduction in the activity of 6-phosphogluconate dehydrogenase was observed in the presence of Al. Changes in enzyme activity were accompanied by changes in levels of G-6-P- the initial substrate in the pentose phosphate pathway. When wheat was exposed for 16 h to a nutrient solution containing aluminium, a 90% reduction in G-6-P concentration was observed. In the Al-tolerant rye cultivar, an increase and subsequently a slight decrease in G-6-P concentration was detected, and after 16 h of Al-stress the concentration of this substrate was still higher than in control plants. This dramatic Al-induced decrease in G-6-P concentration in the Al-sensitive wheat cultivar was associated with a decrease in both the concentration of glucose in the root tips as well as the activity of hexokinase, an enzyme which is responsible for phosphorylation of glucose to G-6-P. However, in the Al-tolerant rye cultivar, the activity of this enzyme remained at the level of control plants during Al-treatment, and the decrease in the concentration of glucose occurred at a much slower rate than in wheat. These results suggest that aluminium ions change cellular metabolism of both wheat and rye root tips. In the Al-sensitive wheat cultivar, irreversible disturbances induced by low doses of Al in the nutrient solution appear very quickly, whereas in the Al-tolerant rye cultivar, cellular metabolism, even under severe stress conditions, is maintained for a long time at a level which allows for root elongation to continue.Abbreviations G-6-PDH glucose-6-phosphate dehydrogenase - 6-PGDH 6-phosphogluconate dehydrogenase - G-6-P glucose-6-phosphate - TEA triethanolamine     

马尾松种源在异质养分环境中的觅养行为差异

选择广东信宜、福建武平、广西岑溪3个不同磷效率特性的马尾松种源,构建同质和异质两种养分环境开展盆栽实验,研究马尾松搜寻利用异质分布养分的获取机制及不同种源觅养行为差异。结果表明,与同质营养环境相比,异质营养环境中马尾松种源具有较高的苗高、地径生长量、较强光合速率和干物质生产能力。研究证实了根系形态可塑性和生理可塑性在马尾松获取异质分布养分中的重要性。马尾松可通过在富养斑块中须侧根的大量增生、对N、P、K等元素的有效吸收提高其觅养能力。马尾松在拓殖富养斑块的初期主要依靠新生侧根的增加和侧根的延长,在拓殖一段时期之后则主要靠新生侧根的生成和须根数量、须根密度的增加来搜寻异质分布养分。异质养分环境中的根系具有较高的养分吸收效率主要缘由在富养斑块中对N、P、K大量的吸收。综合比较分析认为,在3个参试种源中广西岑溪和福建武平种源在异质营养环境中拓殖富养斑块和觅养能力较强,广东信宜种源拓殖和觅养能力相对较弱。     

Localized supply of phosphorus induces root morphological and architectural changes of rice in split and stratified soil cultures

A localized supply of phosphorus may affect root morphology and architecture, and thereby affect phosphorus uptake by rice plants. In the present study, we attempted to test this hypothesis using two rice cultivars representing upland and lowland ecotypes grown in specially designed split and stratified soil cultures with a low-phosphorus red soil. Our data indicate that a localized supply of phosphorus increased both total root length and root fineness, particularly in the high-phosphorus zone. In split culture, plants roots tended to preferentially grow on the high-phosphorus zone, with about 70–75% of the total root length allocated to the high-phosphorus compartment. The total root length on the high-phosphorus side in the split-phosphorus treatment was significantly longer than that in the homogenously high-phosphorus treatment, implying that a phosphorus-deficiency signal from the low-phosphorus side may stimulate the growth of the roots located in the high-phosphorus zone. In stratified soil culture, changes in root morphology and architecture were also observed as indicated by increased total root length, root fineness and relative root allocation in the high-phosphorus layers, again suggesting altered root morphology and preferential root proliferation in the high-phosphorus regions. The induced changes in root morphology and architecture by localized phosphorus supply may have both physiological significance and practical implications in that plants can meet the demand for phosphorus with parts of the roots reaching the high-phosphorus zone, hence localized fertilization methods such as side dressing or banded application of phosphorus fertilizers may both minimize phosphorus fixation by the soil and increase phosphorus uptake efficiency from the fertilizers.     

毛竹种群向针阔林扩张的根系形态可塑性

为了弄清毛竹(Phyllostachys edulis)向针阔林扩张过程中根系的形态可塑性反应,在浙江天目山自然保护区毛竹向针阔林扩张的典型过渡地带,连续区域上设置毛竹纯林、针阔-毛竹混交林(以下简称过渡林)、针阔林3种样地。用根钻法采集样地毛竹根系、针阔树根系并比对其生物量密度、细根比根长、相邻同级侧根节点距等形态特征参数变化。结果表明:随着毛竹的扩张程度增加,林内根系生物量密度增加;且与针阔树竞争过程中毛竹将更多的根系放置于表层;同时在水平方向上随离样株距离的增加未出现明显变化,而针阔树根系则随离样木距离的增加而逐渐减少;毛竹根系比根长明显增加,平均增幅15%;一、二级侧根节点距则均有所下降,毛竹侧根数量增多。这些结果表明毛竹种群可通过根系生物量密度、细根比根长、相邻同级侧根节点距等形态可塑性方式实现向周边森林扩张。