Growth,biomass allocation,and autofragmentation responses to root and shoot competition in Myriophyllum spicatum as a function of sediment nutrient supply

Plant growth, biomass allocation and autofragmentation were investigated in response to root and shoot competition in the submersed macrophyte Myriophyllum spicatum L. growing in two sediment environments. Biomass accumulation and allocation were significantly affected by sediment fertility, with a higher total biomass observed in fertile sediment (average: 4.69 g per plant vs. 1.12 g per plant in infertile sediment). Root-to-shoot ratios were 0.34 and 0.06 in the infertile and fertile sediments, respectively, reflecting the high investment placed on roots under infertile conditions. In the presence of root, shoot, and full competition, whole plant biomass decreased by 18%, 12% and 24% in the infertile sediments, and 23%, 25% and 33% in the fertile sediments, respectively. Root weight ratios (RWRs) increased with root competition by 38% (P < 0.001) and 12% (P = 0.002), while leaf weight ratios (LWRs) decreased with shoot competition by 6% (P = 0.042) and 5% (P = 0.001) in the infertile and fertile sediments, respectively. A total of 406 autofragments were harvested in the fertile sediments, but none were obtained from the infertile sediments. In the control, autofragment number and biomass was 166% and 175% higher compared to the competition treatment. Root and shoot competition resulted in a 21% (P = 0.043) and 18% (P = 0.098) decrease in the autofragment biomass, respectively. These results indicated that M. spicatum responds to different sediment fertility by changing its allocation patterns. Moreover, both root and shoot competition influenced plant growth and autofragmentation, while sediment nutrient availability played an important role in M. spicatum autofragmentation.     

Response in root morphology and nutrient contents of Myriophyllum spicatum to sediment type

Sediment may play an important role during the submerged macrophyte decline in the eutrophication progress. In order to investigate the response in root morphology and nutrient contents of submerged macrophytes Myriophyllum spicatum to sediment, five sediment types were treated and used (five types of sediment were used in the experiment: treatment 1 was nature sediment + sand, a 50:50 (v/v) mixture, treatment 2 was the studied sediment only, treatment 3 was sediment + nitrogen (N, NH₄Cl 400 mg kg^?1), treatment 4 was sediment + phosphorus (P, NaH₂PO₄ 300 mg kg^?1); treatment 5 was sediment + phosphorus (P, NaH₂PO₄ 600 mg kg^?1)). The results show that the root N content was only significantly affected by adding N in sediments and P was elevated by adding N and P. The root mass and its percentage increased at first, the peak values were reached at 35 d, and then decreased. The root growth was restrained by adding sand and N in sediments, root senescence process was delayed at the later experimental time by adding P in sediments. The increase of root volume showed a similar trend to that of root growth, except for plant with P addition where root volume remained high after 35 d. The root volume decreased while the main root number increased significantly by adding sand in sediments. The mean root length and main root diameter were reduced by adding P in sediments. The compatible sediment nutrient condition is necessary to restore submerged macrophytes in a degraded shallow lake ecosystem, and the effect of sediment on the root morphology and nutrient content is one of the important aspects restricting the restoration of submerged macrophytes.     

外源NH_4~+对穗花狐尾藻根系形态和养分吸收的影响

在温室内进行静态实验,以原沉积物(CK)和分别添加0.24%和0.48%氯化铵(SN1和SN2)的沉积物作为底质培养沉水植物,探讨了高NH4+环境中穗花狐尾藻的根系形态特征及营养物质积累与干物质分配策略。结果表明,穗花狐尾藻主根直径(0.432-0.518mm)与主根/侧根比率(1.50-4.39)均随着沉积物NH4+含量升高呈现增大趋势,但单株主根总长(31.64-171.67cm)则在高NH4+环境中显著变短;其中,SN2处理中穗花狐尾藻主根数量(8.17条/株)显著低于SN1(14.67条/株)和CK(14.33条/株)处理,而SN1与CK处理之间差异不显著。SN1和SN2处理中穗花狐尾藻植株全氮含量(55.98和55.19mg/g)均显著高于CK(42.89mg/g)处理,而SN1和SN2处理中穗花狐尾藻植株全磷含量(1.63和1.53mg/g)则比CK处理中(3.71mg/g)显著降低。穗花狐尾藻植株干物质积累量(168.17-405.81mg/株)和全磷积累量(0.25-1.51mg/株)均随沉积物NH4+含量升高而显著下降,但植株全氮积累量(9.12-21.08mg/株)则表现为:SN1CKSN2。全氮和全磷在叶片中的分配率都随着沉积物NH+含量升高而显著降低,而在根系和茎秆中分配率则显著增加。     

Morphological and physiological responses to sediment type and light availability in roots of the submerged plant Myriophyllum spicatum

BACKGROUND AND AIMS: Both sediment and light are essential factors regulating the growth of submerged macrophytes, but the role of these two factors in regulating root morphology and physiology is far from clear. The responses of root morphology and physiology to sediment type and light availability in the submerged plant Myriophyllum spicatum were studied and the hypothesis was tested that a trade-off exists in root growth strategy between internal aeration and nutrient acquisition. METHODS: Plants were grown on two types of sediment (fertile mud and an infertile mixture of mud and sandy loam) and under three levels of light availability (600, 80 and 20 micro mol m(-2) s(-1)) in a greenhouse. KEY RESULTS: The significantly higher alcohol dehydrogenase (ADH) activity in root tissues indicated that oxygen deficiency existed in the plants growing in fertile mud and low (or high) light environments. Significantly, low plant N and P concentrations indicated that nutrient deficiency existed in the mixed sediment and high light environment. As a response to anoxia, plants did not change the porosity of the main roots. The effect of sediment type on root morphology was insignificant under higher light environments, whereas root diameter generally decreased but specific root length (SRL) increased with decreasing light availability. Both low light and fertile mud jointly led to lack of second-order laterals. More biomass was allocated to lateral roots in infertile environments, whereas mass fractions of laterals were lower in low light and mud environments. CONCLUSIONS: These data indicate that this plant can achieve the trade-off between internal aeration and nutrient acquisition by adjusting the structure of the root system and the pattern of biomass allocation to different root orders rather than root morphology and root porosity.     

Morphological responses to water-level fluctuations of two submerged macrophytes,Myriophyllum spicatum and Hydrilla verticillata

Aims Myriophyllum spicatum and Hydrilla verticillata are common submerged macrophytes in the Yangtze River basin. To investigate their tolerances and adaptations to water-level fluctuations, an experiment was conducted in a pond.Methods We designed five different amplitudes of water-level fluctuations as static, ±15, ±30, ±60 and ±90 cm during the 74 days of the experiment. In each amplitude treatment, two cultivation methods were examined as monoculture and mixed culture.Important findings The results showed that M. spicatum had greater morphological responses to water-level fluctuations than H. verticillata. Fluctuating amplitude had significant effects on branch number, shoot length and root dry weight (DW) of M. spicatum, whereas it only had significant effect on branch number of H. verticillata. Both fluctuating amplitude and cultivation method had significant effects on total DW of M. spicatum, which was higher in monoculture than mixed culture. The total DW of H. verticillata was affected by fluctuating amplitude only, and the largest biomass was in the amplitude of ±30 cm. Fruit DW of M. spicatum was largest in the amplitude of ±30 cm, high amplitude of water-level fluctuations would inhibit flowering and seed production. Root DW proportion was significantly affected by fluctuating amplitude and cultivation method in both species, and the root DW proportion of M. spicatum was significantly higher in the amplitude-of ±90 cm. We conclude that moderate amplitude of water-level fluctuations can promote the distribution and growth of both species, and in order to accelerate the restoration of both species in natural habitats, the optimum amplitude should be keep at ±15 to ±30 cm.     

Growth and shoot: root ratio of seedlings in relation to nutrient availability

The influence of mineral nutrient availability, light intensity and CO₂ on growth and shoot:root ratio in young plants is reviewed. Special emphasis in this evaluation is given to data from laboratory experiments with small Betula pendula plants, in which the concept of steady-state nutrition has been applied.Three distinctly different dry matter allocation patterns were observed when growth was limited by the availability of mineral nutrients: 1, Root growth was favoured when N, P or S were the major growth constraints. 2, The opposite pattern obtained when K, Mg and Mn restricted growth. 3, Shortage of Ca, Fe and Zn had almost no effect on the shoot:root ratio. The light regime had no effect on dry matter allocation except at very low photon flux densities (< 6.5 mol m^-2 day^-1), in which a small decrease in the root fraction was observed. Shortage of CO₂, on the other hand, strongly decreased root development, while an increase of the atmospheric CO₂ concentration had no influence on dry matter partitioning. An increased allocation of dry matter to below-ground parts was associated with an increased amount of starch in the tissues. Depletion of the carbohydrate stores occurred under all conditions in which root development was inhibited. It is concluded that the internal balance between labile nitrogen and carbon in the root and the shoot system determines how dry matter is being partitioned in the plant. The consistency of this statement with literature data and existing models for shoot:root regulation is examined.     

根茎型植物扁秆荆三棱对光照强度和养分水平的生长响应及资源分配策略

自然界中光照和养分因子常存在时空变化,对植物造成选择压力。克隆植物可通过克隆生长和生物量分配的可塑性来适应环境变化。尽管一些研究关注了克隆植物对光照和养分因子的生长响应,但尚未深入全面了解克隆植物对光照和养分资源投资的分配策略。以根茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis)为研究对象,在温室实验中,将其独立分株种植于由2种光照强度(光照和遮阴)和4种养分水平(对照、低养分、中养分和高养分)交叉组成的8种处理组合中,研究了光照和养分对其生长繁殖及资源贮存策略的影响。结果表明,扁秆荆三棱的生长、无性繁殖及资源贮存性状均受到光照强度的显著影响,在遮阴条件下各生长繁殖性状指标被抑制。且构件的数目、长度等特征对养分差异的可塑性响应先于其生物量积累特征。在光照条件下,高养分处理的总生物量、叶片数、总根茎分株数、长根茎分株数、总根茎长、芽长度、芽数量等指标大于其他养分处理,而在遮阴条件下,其在不同养分处理间无显著差异,表明光照条件可影响养分对扁秆荆三棱可塑性的作用,且高营养水平不能补偿由于光照不足而导致的生长能力下降。光照强度显著影响了总根茎、总球茎及大、中、小球茎的生物量分配,遮阴条件下,总生物量减少了对地下部分根茎和球茎的分配,并将有限的生物量优先分配给小球茎。总根茎的生物量分配未对养分发生可塑性反应,而随着养分增加,总球茎分配下降,说明在养分受限的环境中球茎的贮存功能可缓冲资源缺乏对植物生长的影响。在相同条件下,根茎生物量对长根茎的分配显著大于短根茎,以保持较高的繁殖能力;而总球茎对有分株球茎的生物量分配小于无分株球茎,表明扁秆荆三棱总球茎对贮存功能的分配优先于繁殖功能。研究为进一步理解根茎型克隆植物对光强及基质养分环境变化的生态适应提供了依据。     

Growth and shoot:root partitioning of spinach plants as affected by nitrogen supply

Spinach plants (Spinacia oleracea L.) were grown hydroponically in fixed environmental conditions either at full nitrate availability (11·8mol m^-3) or at a suboptimum relative nitrate addition rate of 0·20d^-1, 0·15d^-1 or 0·10d^-1 respectively, the other nutrients being adequately provided. The relative growth rate (RGR) of the plants varied significantly with the nutrition treatment and decreased during development in all treatments. The concentration of reduced nitrogen in the plants grown at full nitrate availability did not change significantly during the experimental growth period and nitrate accumulation was substantial. After an adaptation period, the concentration of reduced nitrogen in the plants at the suboptimum nitrate addition rates increased during growth and was lowest at the lowest relative nitrate addition rate. Nitrate uptake was almost complete in the suboptimum treatments and nitrate accumulation was negligible as long as the concentration of reduced nitrogen was below 2·0 mmol (g dry weight)^-1. The RGR of all plants was proportional to the concentration of reduced nitrogen in the plant minus a minimal tissue concentration required for growth. However, the proportionality factor was inversely related to the plant mass. This relationship was summarized in an empirical model which explained 98·7% of the variance of the dry weight (log scale) data of all treatments at all harvests. The model was compared with other growth models found in the literature. The shoot/root weight ratio increased from 2 to 4 if nitrate provision was not limiting, and initially, this ratio decreased at suboptimum nitrate provision but increased at higher growth stages. Possible explanations of the dynamics of dry matter partitioning are discussed in relation to models.     

Photosynthetic and biomass allocation responses of Liquidambar styraciflua (Hamamelidaceae) to vine competition

This 2-year field study examined stomatal conductance, photosynthesis, and biomass allocation of Liquidambar styraciflua saplings in response to below- and aboveground competition with the vines Lonicera japonica and Parthenocissus quinquefolia. Vine competition did not affect stomatal conductance of the host trees. The leaf photosynthetic capacity and photosynthetic nitrogen-use efficiency were significantly reduced by root competition with vines, either singly or in combination with aboveground competition, early in the second growing season. However, such differences disappeared by the end of the second growing season. Trees competing below ground with vines also had lower allocation to leaves compared with steins. Aboveground competition with vines resulted in reduced photosynthetic capacity per unit leaf area, but not per unit leaf weight, in trees. No correlation was found between single leaf photosynthetic capacity and tree growth. In contrast, a high positive correlation existed between allocation to leaves and diameter growth. Results from this study suggest that allocation patterns are more affected than leaf photosynthesis in trees competing with vines.     

Influence of Myriophyllum spicatum L. on the species composition,biomass and primary productivity of phytoplankton

Myriophyllum spicatum L. in enclosures initially increased phytoplankton productivity per unit biomass relative to enclosures from which it was absent. Phytoflagellates prevailed during the entire season in the presence of M spicatum, while desmids were dominant in the other communities.     

光周期对穗花狐尾藻生长、开花与种子形成的影响

通过人工补光和遮光处理,初步研究了光周期对狐尾藻生长和开花的影响。研究发现:8h短日照条件不利于狐尾藻的生长和花序形成,导致狐尾藻不能正常开花和结实;与自然日照长度(平均日照14h)条件下相比,16h长光照条件可以增加植株的高度和分枝数,形成更大的生物量;但24h全光照条件对生长有抑制作用。长光照条件下,狐尾藻花序形成时间和开花时间均比自然条件下延迟,形成花序的数目也显著较自然条件下的少,但长光照条件下形成的种子比自然条件下形成的种子具有更高的萌发率。     

Plant allocation of carbon to defense as a function of herbivory,light and nutrient availability

We use modeling to determine the optimal relative plant carbon allocations between foliage, fine roots, anti-herbivore defense, and reproduction to maximize reproductive output. The model treats these plant components and the herbivore compartment as variables. Herbivory is assumed to be purely folivory. Key external factors include nutrient availability, degree of shading, and intensity of herbivory. Three alternative functional responses are used for herbivory, two of which are variations on donor-dependent herbivore (models 1a and 1b) and one of which is a Lotka–Volterra type of interaction (model 2). All three were modified to include the negative effect of chemical defenses on the herbivore. Analysis showed that, for all three models, two stable equilibria could occur, which differs from most common functional responses when no plant defense component is included. Optimal strategies of carbon allocation were defined as the maximum biomass of reproductive propagules produced per unit time, and found to vary with changes in external factors. Increased intensity of herbivory always led to an increase in the fractional allocation of carbon to defense. Decreases in available limiting nutrient generally led to increasing importance of defense. Decreases in available light had little effect on defense but led to increased allocation to foliage. Decreases in limiting nutrient and available light led to decreases in allocation to reproduction in models 1a and 1b but not model 2. Increases in allocation to plant defense were usually accompanied by shifts in carbon allocation away from fine roots, possibly because higher plant defense reduced the loss of nutrients to herbivory.     

Spatial heterogeneity in soil nutrient supply modulates nutrient and biomass responses to multiple global change drivers in model grassland communities

Changes in the atmospheric concentration of carbon dioxide ([CO₂]), nutrient availability and biotic diversity are three major drivers of the ongoing global change impacting terrestrial ecosystems worldwide. While it is well established that soil nutrient heterogeneity exerts a strong influence on the development of plant individuals and communities, it is virtually unknown how nutrient heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. We conducted a microcosm experiment to evaluate the effect of simultaneous changes in [CO₂], nutrient heterogeneity (NH), nutrient availability (NA) and species evenness on the biomass and nutrient uptake patterns of assemblages formed by Lolium perenne, Plantago lanceolata and Holcus lanatus. When the nutrients were heterogeneously supplied, assemblages exhibited precise root foraging patterns, and had higher above‐ and belowground biomass (average increases of 32% and 29% for above‐ and belowground biomass, respectively). Nutrient heterogeneity also modulated the effects of NA on biomass production, complementarity in nitrogen uptake and below: aboveground ratio, as well as those of [CO₂] on the nutrient use efficiency at the assemblage level. Our results show that nutrient heterogeneity has the potential to influence the response of plant assemblages to simultaneous changes in [CO₂], nutrient availability and biotic diversity, and suggest that it is an important environmental factor to interpret and assess plant assemblage responses to global change.     

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     

Fertile sediment and ammonium enrichment decrease the growth and biomechanical strength of submersed macrophyte Myriophyllum spicatum in an experiment

Decline of submersed macrophytes has occurred in eutrophic lakes worldwide. Little is known about effects of nutrient enrichment on biomechanical properties of submersed macrophytes. In a 30-day experiment, Myriophyllum spicatum was cultured in aquaria containing two types of sediment (mesotrophic clay vs. fertile loam) with contrasting water NH₄ ⁺ concentrations (0 vs. 3.0 mg L^?1 NH₄–N). The plant growth, shoot and root morphology, stem biomechanical properties, and stem total nonstructure carbohydrates content (TNC) were examined. The NH₄ ⁺-enriched water, particularly combined with the fertile sediment, caused adverse effects on M. spicatum as indicated by reductions in the growth, stem biomechanical properties (tensile force, bending force and structural stiffness), and TNC content. These results indicate that increased sediment fertility and water NH₄ ⁺-enrichment made the plant more fragile and vulnerable to hydraulic damage, particularly for the upper stem, implying that M. spicatum was prone to uprooting and fracture by hydraulic force, and the broken fragment from parent shoot of M. spicatum might have low-survival potential due to its low-TNC content. This may be a mechanical aspect for the decline of submersed macrophytes and makes it more difficult to restore submersed vegetation in the eutrophic lakes.