Inhibition of seedling survival under Rhododendron maximum (Ericaceae): could allelopathy be a cause?

In the southern Appalachian mountains a subcanopy species, Rhododendron maximum, inhibits the establishment and survival of canopy tree seedlings. One of the mechanisms by which seedlings could be inhibited is an allelopathic effect of decomposing litter or leachate from the canopy of R. maximum (R.m.) on seed germination, root elongation, or mycorrhizal colonization. The potential for allelopathy by R.m. was tested with two bioassay species (lettuce and cress), with seeds from four native tree species, and with three ectomycorrhizal fungi. Inhibitory influences of throughfall, fresh litter, and decomposed litter (organic layer) from forest with R.m. (+R.m. sites) were compared to similar extractions made from forest without R.m. (-R.m. sites). Throughfall and leachates of the organic layer from both +R.m. and -R.m. sites stimulated germination of the bioassay species above that of the distilled water control, to a similar extent. There was an inhibitory effect of leachates of litter from +R.m. sites on seed germination and root elongation rate of both bioassay species compared with that of litter from -R.m. sites. Native tree seed stratified in forest floor material from both forest types had a slightly higher seed germination rate compared with the control. A 2-yr study of seed germination and seedling mortality of two tree species, Quercus rubra and Prunus serotina, in field plots showed no significant influence of litter or organic layer from either forest type. Incorporating R.m. leaf material into the growth medium in vitro depressed growth of one ectomycorrhizal species but did not affect two other species. Leaf material from other deciduous tree species depressed ectomycorrhizal growth to a similar or greater extent as leaf material from R.m. In conclusion, R.m. litter can have an allelopathic effect on seed germination and root elongation of bioassay species as well as some ectomycorrhizal species. However, this allelopathic affect is not manifest in field sites and is not likely to be an important cause for the inhibition of seedling survival within thickets of R.m.     

Interactions between vesicular-arbuscular mycorrhizal fungi and asparagus

Summary The vesicular-arbuscular mycorrhizal (VAM) fungus,Glomus versiforme increased significantly the growth ofAsparagus officinalis under controlled conditions using Turface as the growth medium. The growth responses, including increases in root fresh weight, numbers of shoots, shoot dry weight, and shoot height follow a pattern similar to other mycorrhizal systems. Indigenous VAM fungi appeared to have negative effects on average shoot fresh and dry weight, number of shoots per pot and average shoot height on one year oldA. officinalis seedlings obtained from the field and grown under controlled conditions. These results may be due either to the high levels of soluble phosphate present in the soil or the ineffectiveness of the particular indigenous fungi as mycorrhizal fungi in asparagus. Indigenous mycorrhizal fungi overwinter in asparagus root crown as vesicles and as external and internal hyphae. Soil obtained from the same fields as the one year old crowns was a good source of mycorrhizal inoculum for sterile seedlings.     

CONTRASTING GERMINATION AND SEEDLING GROWTH OF BETULA ALLEGHANIENSIS AND RHUS TYPHINA SUBJECTED TO VARIOUS AMOUNTS AND TYPES OF PLANT LITTER

Several characteristics of the environment, such as plant litter, may interact with species characteristics, such as seed size, to determine patterns of plant establishment and thereby influence abundance and spatial location of plants long after the factor has itself disappeared. To determine if litter might differentially affect seedling establishment of Betula alleghaniensis Britton and Rhus typhina L., two woody species with different sized seeds, we conducted two greenhouse experiments. We documented emergence, morphology, and growth patterns of seedlings covered by three amounts each of Fagus grandifolia Ehrh. leaves or Tsuga canadensis L. needles. All amounts of litter significantly reduced and delayed final Betula alleghaniensis seedling emergence, and the resultant seedlings had reduced root: shoot ratios, longer hypocotyls, and were less robust. Litter did not affect Rhus typhina emergence but did alter biomass allocation: stem, leaf, and root biomass of Rhus seedlings was reduced by greater amounts of litter, and by leaf litter relative to needle litter. We attribute the different responses of Betula and Rhus to differences in seed size (Betula mean seed size = 1.0 mg; Rhus mean seed size = 8.5 mg) and germination cues (Betula requires light, Rhus requires chemical or heat scarification), and suggest that differential response to litter may contribute to coexistence among these sympatric species.     

种子散布位置对格氏栲幼苗根系形态及生长特征的影响

种子从母株掉落于地面萌发后,其根系在不同散布位置(凋落物上层、土壤表层和凋落物下层)的生长形态影响幼苗定居及建成,而目前对其根系形态及生长特征的了解并不充分,限制了对幼苗根系在不同散布下适应策略的理解。为此,以格氏栲(Castanopsis kawakamii)为研究对象,通过模拟种子在凋落物中位置,设置凋落物上层(种子下层铺垫2 cm和4 cm凋落物,U2和U4处理)、土壤表层与凋落物下层(种子上层覆盖0、2、4、6 cm和8 cm凋落物,CK、D2、D4、D6和D8处理)等3种散布,探讨不同散布位置对格氏栲幼苗根系9个生长指标的影响。结果表明：(1)种子散布位置对幼苗根干物质质量具有显著影响,D2处理达最大值。(2)D2处理的幼苗根长、根表面积、根尖数、分枝数和比根长高于其它处理;根系平均直径在D6处理达最大值。(3)相关分析表明根长、根表面积、根尖数、分枝数和比根长与根系平均直径呈显著负相关关系。(4)对根系9个生长指标提取主成分后聚类为4个类群,D2与D4处理各划分一类;U2与U4处理划分一类,其余三个处理划分一类。综上所述,凋落物浅层覆盖(D2处理)适宜格氏栲根系生长;凋落物...     

Seed Film Coating with Uniconazole Improves Rape Seedling Growth in Relation to Physiological Changes Under Waterlogging Stress

Waterlogging is an important constraint of global production of rape. The effects of seed film coating with 0.0075% uniconazole on the growth and physiology was investigated using seedlings from three varieties of rape (Brassica napus L.) subjected to waterlogging. While seed coating with uniconazole had no significant effect on germination percentage during waterlogging stress, it enhanced root vigour, increased root length, root volume and root dry weight. It also significantly enhanced leaf dry weight and ratio of root to shoot, but induced a significant decrease in shoot height and stem dry weight. Seed film coating with uniconazole also significantly increased the activities of the antioxidant enzymes, POD, CAT and SOD, and soluble sugar concentration during waterlogging. Thus, it suggests that seed film coating with uniconazole at a suitable concentration can improve rape seedling growth and increase seedling establishment during waterlogging.     

Identification and Phytotoxicity Assessment of Phenolic Compounds in Chrysanthemoides monilifera subsp. monilifera (Boneseed)

Chrysanthemoides monilifera subsp. monilifera (boneseed), a weed of national significance in Australia, threatens indigenous species and crop production through allelopathy. We aimed to identify phenolic compounds produced by boneseed and to assess their phytotoxicity on native species. Phenolic compounds in water and methanol extracts, and in decomposed litter-mediated soil leachate were identified using HPLC, and phytotoxicity of identified phenolics was assessed (repeatedly) through a standard germination bioassay on native Isotoma axillaris. The impact of boneseed litter on native Xerochrysum bracteatum was evaluated using field soil in a greenhouse. Collectively, we found the highest quantity of phenolic compounds in boneseed litter followed by leaf, root and stem. Quantity varied with extraction media. The rank of phenolics concentration in boneseed was in the order of ferulic acid > phloridzin > catechin > p-coumaric acid and they inhibited germination of I. axillaris with the rank of ferulic acid > catechin > phloridzin > p-coumaric acid. Synergistic effects were more severe compared to individual phenolics. The litter-mediated soil leachate (collected after15 days) exhibited strong phytotoxicity to I. axillaris despite the level of phenolic compounds in the decomposed leachate being decreased significantly compared with their initial level. This suggests the presence of other unidentified allelochemicals that individually or synergistically contributed to the phytotoxicity. Further, the dose response phytotoxic impacts exhibited by the boneseed litter-mediated soil to native X. bracteatum in a more naturalistic greenhouse experiment might ensure the potential allelopathy of other chemical compounds in the boneseed invasion. The reduction of leaf relative water content and chlorophyll level in X. bracteatum suggest possible mechanisms underpinning plant growth inhibition caused by boneseed litter allelopathy. The presence of a substantial quantity of free proline in the target species also suggests that the plant was in a stressed condition due to litter allelopathy. These findings are important for better understanding the invasive potential of boneseed and in devising control strategies.     

Comparative greenhouse study of Eucalyptus grandis in vitro plantlets and half-sib seedlings,II. Dry matter accumulation and relative distribution

In vitro directly micropropagated plantlets from three selected five-year-old Eucalyptus grandis Hill ex. Maiden hybrids were compared to their related half-sib seedlings for growth and growth pattern parameters under greenhouse conditions used for operational seedling production. The oven dry weights were determined from stem, leaf, and root samples collected every 40 days for four times. Relative growth rate, net assimilation rates and shoot:root ratio were calculated. Survival was 98% and 95% for plantlets and seedlings, respectively. Significant differences were observed between parents in terms of shoot and root dry weights and their ratios with similar ranking among plantlets and seedlings, suggesting genetic control over these traits. Plantlets started with significantly higher root: shoot ratios and stem, leaf, root, and total dry weight. Although seedlings had higher relative growth and net assimilation rates, all the initial differences decreased sharply over time.     

铁核桃根与叶水浸提液对果园间作绿肥植物的化感作用及其生理机制

以西南地区具有代表性的16种绿肥植物为受体材料,采用培养皿药膜法研究了铁核桃(Juglans sigillata)根水浸提液对受体种子发芽率及幼苗鲜重、干重的化感效应;并进一步研究了铁核桃根、叶水浸提液和胡桃醌对化感效应存在明显差异的4种绿肥植物(绿豆、红三叶、白三叶、花生)种子萌发与幼苗生长以及抗氧化酶特性的影响,以筛选适宜中国西南地区核桃园种植的绿肥植物,探讨核桃根和凋落物对绿肥作物的化感作用机制。结果表明:(1)铁核桃根水浸提液对绿豆的发芽率没有影响,但对绿豆幼苗鲜重和干重有显著抑制作用,而对其他15种绿肥的发芽率和鲜重、干重均有抑制作用。(2)胡桃醌显著抑制绿豆种子萌发,而铁核桃根或叶水浸提液对绿豆种子萌发没有影响。(3)铁核桃根或叶水浸提液以及胡桃醌对绿肥植物幼苗生长的化感效应趋势一致,但核桃根或叶水浸提液的化感效应强于胡桃醌。(4)绿豆幼苗在铁核桃根或叶水浸提液以及胡桃醌处理下,超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)的活性均高于其他3种(红三叶、白三叶、花生)受体幼苗,表明绿豆清除活性氧能力高,细胞受损害程度较低,受化感作用影响最弱。研究认为,绿豆为适宜中国西南地区幼龄核桃园种植的间作绿肥植物。     

Allelopathic potential of Bothriochloa laguroides var. laguroides (DC.) Herter (Poaceae: Andropogoneae)

Bothriochloa laguroides var. laguroides (DC.) Herter, a native bluestem of America, has been shown to produce many biologically active compounds. The allelopathic potential of aqueous extracts from roots, stems and leaves was examined. Lettuce seeds (Lactuca sativa) and maize (Zea mays), the common allelopathy bioassay systems, as well as seeds from two native species, wintergreen paspalum (Paspalum guenoarum) and lovegrass (Eragrostis curvula), were germinated in the presence of aqueous extracts. Percent seed germination, root and shoot elongation were measured. After 4 and 7 days root, stem and leaf extracts caused inhibition of root and shoot elongation in all four species tested. Aqueous extracts were generally less inhibitory to seed germination. Aqueous extracts from different parts of B. laguroides var. laguroides show therefore allelopathic effects inhibiting, in particular, growth of competing plants.     

侧柏不同器官水提取物对油松种子萌发和幼苗生长的他感效应

研究了侧柏（Platycladus orientalis）根、枝、叶、果实不同浓度水提取物及枝叶挥发油对油松种子萌发和幼苗生长的他感效应。结果表明：侧柏各器官及挥发油都含有化感物质,均对油松种子萌发及幼苗生长表现出“高抑低促”效应,而且低浓度时的促进作用明显强于高浓度时的抑制作用。与对照相比,在浓度为2 mg·mL^-1时,侧柏根水提取物对油松幼苗根长和鲜重的促进能力最强,其根长和鲜重分别比对照提高了79.78%和376.60%;侧柏叶水提取物对油松幼苗苗高促进能力最强,比对照提高了102.41%;侧柏枝水提取物对油松种子发芽的促进能力最强,发芽率比对照提高了65.63%。而在浓度为30 mg·mL^-1时,除了侧柏根和枝水提取物对油松种子萌发（87.50%）和鲜重（85.11%）表现出明显的抑制作用外,其它提取物对油松幼苗根长和苗高生长的抑制作用均不明显。侧柏枝叶挥发油对油松种子萌发,幼苗生长均表现为促进作用,特别对油松种子萌发具有显著的促进作用,其促进率达70.37%。说明侧柏和油松混交,可能有利于油松树木生长及生产力的提高。     

Effects of artificial and western spruce budworm (Lepidoptera: Tortricidae) defoliation on growth and biomass allocation of Douglas-fir seedlings

Artificial defoliation has been used commonly to simulate defoliation by insect herbivores in experiments, in spite of the fact that obvious differences exist between clipping foliage and natural defoliation due to insect feeding. We used a greenhouse experiment to compare the effects of artificial and western spruce budworm (Choristoneura occidentalis Freeman) defoliation on the growth and biomass allocation of 3-yr old half-sib seedlings from mature Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco variety glauca] trees that showed phenotypic resistance versus susceptibility to budworm defoliation in the forest. Artificial clipping of buds mimicked the effects of budworm feeding on total seedling biomass when 50% of the terminal buds were damaged. However, artificial defoliation decreased seedling height, relative growth rate of height, and shoot: root ratio more than budworm defoliation, whereas budworm defoliation decreased stem diameter relative growth rate more than artificial defoliation. Half-sib seedling progeny from resistant maternal tree phenotypes had greater height, diameter, biomass, and shoot: root ratio than seedlings from susceptible phenotypes. We concluded that careful artificial defoliation could generally simulate effects of budworm defoliation on total biomass of Douglas-fir seedlings, but that the two defoliation types did not have equal effects on biomass allocation between shoot and root. Further, an inherently higher growth rate and a greater allocation of biomass to shoot versus root are associated with resistance of Douglas-fir trees to western spruce budworm defoliation.     

Regulation of isoflavone production in hydroponically grown Pueraria montana (kudzu) by cork pieces, XAD-4, and methyl jasmonate

A mini-hydroponic growing system was employed for seedlings of kudzu vine (Pueraria montana) and contents of isoflavones (daidzein, genistein, daidzin, genistin, and puerarin) from shoot and root parts of seedlings were analyzed quantitatively. In addition, exogenous cork pieces, polymeric adsorbent, XAD-4, and universal elicitor, methyl jasmonate (MeJA), were used to regulate the production of these isoflavones. It was shown that cork pieces up-regulate the production of daidzein and genistein up to seven- and eight-fold greater than the levels obtained for control roots. In contrast, levels of glucosyl conjugates, daidzin and genistin, decrease up to five- and eight-fold, respectively. Cork treatment also induces the excretion of the root isoflavone constituents into the growth medium. Minimal levels of isoflavones are absorbed by the cork pieces. XAD-4 stimulates the production of glucosyl conjugates, daidzin and genistin, in root parts about 1.5-fold greater than that obtained in control roots. These are the highest amounts of daidzin and genistin that are observed (5.101 and 6.759 mg g⁻¹ dry weight, respectively). In contrast to these two adsorbents, MeJA increases the accumulation of isoflavones in shoot rather than in root parts of seedlings, about three- to four-fold over control levels, with the exception of genistein. These studies reveal new observations on the regulation of isoflavone production in hydroponically grown Pueraria montana plants by two adsorbents (cork pieces and XAD-4) and MeJA elicitor.     

本氏针茅根际土提取物对本氏针茅及其优势种种子萌发的影响

以本氏针茅及其优势种大针茅和百里香种子为受体,采用生物检测的方法研究了0(对照)、0.1、0.2和0.5g/mL本氏针茅根际土甲醇浸提液和水浸提液对3种受体种子萌发的影响,以探究草地演替过程中物种间的生态关系和化感作用的表现形式。结果显示:(1)甲醇浸提液对3种受体种子的发芽均有抑制作用;水浸提液对本氏针茅发芽有促进作用,对百里香有部分促进作用,而对大针茅有抑制作用。(2)甲醇浸提液对本氏针茅和大针茅胚根和胚芽生长无显著影响,而抑制百里香胚根和胚芽的生长;水浸提液对本氏针茅胚根生长无显著影响,对本氏针茅胚芽生长具有显著促进作用,而对其它两种受体胚根和胚芽均无显著影响。(3)各种浓度甲醇浸提液对大针茅和百里香生长表现出了化感综合抑制效应,对本氏针茅生长于0.2g/mL时表现出化感综合抑制效应,其它2种浓度(0.1、0.5g/mL)表现为化感综合促进效应;水浸提液对本氏针茅和百里香表现为化感综合促进效应,而对大针茅表现为化感综合抑制效应。研究表明,本氏针茅根际土甲醇浸提液和水浸提液的化感物质存在差异;不同受体对同一浸提液反应不同,同一受体对不同浸提液的反应也不同。     

Litter Quantity and Nitrogen Immobilization Cause Oscillations in Productivity of Wild Rice (Zizania palustris L.) in Northern Minnesota

Microbial immobilization of nitrogen (N) in litter from one year’s production may cause oscillations in biomass production if it delays N availability the following year. We tested whether shoot and root litter and plant density affect biomass and seed production of populations of wild rice (Zizannia palustris L.) grown in 378 l stock tank mesocosms over four consecutive years. Half the tanks were thinned to a uniform seedling density whereas density in the remaining tanks was allowed to fluctuate ad libitum. Litter treatments included both shoot litter removal, leaving only root litter, and retaining shoot litter intact with root litter. A separate greenhouse fertilizer experiment tested whether N and/or phosphorus (P) limited productivity. Responses to N additions were much stronger than to P additions. Annual production and N availability in the tanks were correlated with each other and followed a concurrent cycle from 2004 to 2008. Furthermore, production in tanks with shoot + root litter did not fluctuate more than tanks with only root litter. Root litter immobilized more nitrogen and for a longer period than shoot litter. Neither litter immobilized P. Density did not affect mean seed weight, total seed production, or mean plant weight, but total seed production declined in years following productive years and was high only following years of low litter production. Root litter may therefore be primarily responsible for the delays in N availability that cause cycles in biomass and seed production. Consequently, both wild rice litter quantity and quality play central roles in production and population dynamics of wild rice stands.     

Metabolism of phenolic compounds in <Emphasis Type="Italic">Vitis riparia</Emphasis> seeds during stratification and during germination under optimal and low temperature stress conditions

We studied the alterations in phenolic compounds in grape seeds during their stratification and germination under optimal conditions (+25 °C) and at low temperature (+10 °C). Biological materials in the study were seeds of Vitis riparia. Phenolic compounds were extracted from defatted seeds using 80 % methanol or 80 % acetone. The content of total phenolics was determined with the Folin-Ciocalteau reagent, while the content of tannins was determined by vanillin assay and the protein (BSA) precipitation method. The RP-HPLC method was used to determine phenolic compounds (phenolic acids, catechins) in the extracts. High amounts of tannins, catechins, gallic acid and lesser amounts of p-coumaric acid were found in the seeds. The content of total phenolics in acetone extracts was higher than that obtained using methanol. The amounts of phenolic acids and tannins found in V. riparia seeds after stratification were much lower. It may confirm a possible role of these compounds in dormancy of V. riparia seeds. After 72 h of low temperature treatment, inhibition of grape root growth and biochemical changes in seeds were detected. The chilling stimulated increased accumulation of some phenolic compounds (free gallic acid and catechins) in the seeds. These substances can protect plants against some abiotic stressors.     

Chemical,physical and biological control of carrot seedling diseases

In naturally infested soil containingPythium ultimum, P. acanthicum andPhytophthora megasperma, onlyP. ultimum was associated with root rot and damped-off seedlings. Damping-off was promoted by low soil temperatures and by flooding. Seedling stands were markedly reduced when seed was pre-incubated in soil at 12°C but not at 25°C or 35°C. Dusting carrot seed with metalaxyl significantly increased seedling stands in the field at rates from 1.5–6 g kg⁻¹ seed and in both flooded and unflooded, naturally infested soil at 3.15 g kg⁻¹. In greenhouse experiments using artifically infested soil,P. ultimum andP. paroecandrum caused damping-off of carrot seedlings andRhizoctonia solani reduced root and shoot weights.R. solani caused damping-off in nutrient-enriched soil.P. acanthicum andP. megasperma were not pathogenic to seedlings, although both fungi colonized roots. Soil populations of allPythium spp., particularlyP. ultimum, increased during growth of seedlings and population growth ofP. megasperma was promoted by periodic flooding. Infestation of soil withP. acanthicum did not reduce damping-off of carrot seedlings byP. ultimum orP. paroecandrum, but significantly increased root and shoot weights and decreased root colonization byR. solani P. acanthicum has potential as a biocontrol agent againstR. solani.     

冷蒿对三种禾本科植物种子萌发和幼苗生长的化感作用

研究了冷蒿茎叶水浸提液及冷蒿斑块土壤对羊草草原3种禾本科植物(羊草、克氏针茅和糙隐子草)种子萌发及幼苗生长的影响.结果表明:冷蒿茎叶水浸提液使受试植物种子的发芽指数降低,平均发芽时间延长.羊草、克氏针茅和糙隐子草幼苗苗高及克氏针茅幼苗根长均受冷蒿茎叶水浸提液抑制;浸提液浓度≥0.075g.ml-1时显著抑制羊草幼苗根长生长,≤0.05g.ml-1时则无显著影响;而茎叶水浸提液对糙隐子草幼苗根长有"低促高抑"的作用.冷蒿斑块土壤抑制受试植物幼苗生长;3种植物对冷蒿化感效应的敏感性依次为克氏针茅羊草糙隐子草,且幼苗根长的敏感性均大于苗高.     

Rapid bioassays to evaluate the plant growth promoting activity of <Emphasis Type="Italic">Ascophyllum nodosum</Emphasis> (L.) Le Jol. using a model plant, <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> (L.) Heynh

Ascophyllum nodosum extract products are used commercially in the form of liquid concentrate and soluble powder. These formulations are manufactured from seaweeds that are harvested from natural habitats with inherent environmental variability. The seaweeds by themselves are at different stages of their development life-cycle. Owing to these differences, there could be variability in chemical composition that could in turn affect product consistency and performance. Here, we have tested the applicability of using Arabidopsis thaliana as a model to study the activity of two different extracts from A. nodosum. Three different bioassays: Arabidopsis root-tip elongation bioassay, Arabidopsis liquid growth bioassay and greenhouse growth bioassay were evaluated as growth assays. Our results indicate that both extracts promoted root and shoot growth in comparison to controls. Further, using Arabidopsis plants with a DR5:GUS reporter gene construct, we provide evidence that components of the commercial A. nodosum extracts modulates the concentration and localisation of auxins which could account, at least in part, for the enhanced plant growth. The results suggest that A. thaliana could be used effectively as a rapid means to test the bioactivity of seaweed extracts and fractions.     

芒萁对几种杂草和农作物的生化他感作用

本文研究了芒萁植株地上部分和地下部分水提液对几种杂草和农作物的生化他感作用。证明：（１） 芒萁水提液对供试杂草及白菜、萝卜的种子萌发有显著或极显著抑制作用,对全部供试幼苗的根生长和多数供试幼苗的苗高有显著抑制作用;（２） 地上部分的生化他感作用强于地下部分;（３） 芒萁对供试植物幼苗根生长的抑制作用强于对茎、叶的作用     

岩生植物金发草对作物的化感效应

采用生物测定法研究金发草根、茎、叶水浸提液对水稻、小麦、油菜和莴苣等4种受体植物的化感作用。结果表明,金发草根、茎、叶水浸提液对4种受体植物种子萌发和幼苗生长均具化感作用,且随着水浸提液浓度的增高而增强。除水稻外,小麦、油菜、莴苣的发芽速率均受金发草水浸提液的显著抑制,但对最终发芽率影响不显著,说明金发草对植物种子萌发的化感作用主要是延迟种子发芽时间;金发草对作物幼苗生长的化感作用总体上表现为促进地上部分生长而抑制地下部分的生长。受体植物不同发育期对金发草化感作用的敏感程度不同,幼苗期高于萌发期;金发草叶的化感作用最强,其综合效应分别为根与茎的2.14倍和2.56倍。