Influence of soil temperature during root appearance on the trajectory of nodal roots of field grown maize

Trajectories of maize nodal roots were studied to test the hypothesis that roots which appear on a common internode have similar geometrical characteristics, and to assess the effect of soil temperature on root trajectory. Treatments consisted of three sowing dates, a comparison between mulched and non-mulched soil, a replication of one sowing date for two years in two locations, and a comparison between two cultivars at one sowing date. All these sources of variation, except the cultivar, had an appreciable effect on the trajectories of roots which appeared on the first four internodes. The horizontal component of the trajectory differed significantly between treatments, ranging from 93 to 700 mm in roots which appeared on the second internode, and from 71 to 569 for those on the third internode. The original hypothesis had, therefore, to be rejected. Mean soil temperature during the 100°C.days after root appearance accounted for the differences in trajectory between location, year, sowing date and mulch treatments, and for the differences between internodes within each location. The critical period during which temperature affected root trajectory probably began at root appearance, and ended between 50 and 100°C days after root appearance, i.e. when the root was less than 100 mm long.     

柠条人工林细根不同分枝根序寿命估计

植物细根在发育结构上表现的形态特征和生理功能异质性影响细根寿命的准确估计,因此了解分枝根序细根寿命差异对于深入认识细根的周转过程和陆地生态系统碳分配具有重要意义。采用微根管(Minirhizotron)技术对晋西北黄土区的五年生柠条(Caragana Korshinskii Kom.)人工林细根的生长过程进行了为期3a(2007—2009年)的追踪观测,分析了不同因素(土层深度、季节变化、空间位置)对一级根和高级根寿命的影响。结果表明:(1)在各土层深度处,一级根的中值寿命均低于高级根中值寿命,其中一级根中值寿命表现随土层深度增加而增加趋势,而高级根除表层0—20 cm中值寿命较短外,各土层间变化趋势不明显,40—60 cm、80—100 cm土层高级根在观测期结束时其累积存活率仍在50%以上;(2)不同季节出生一级根和高级根的中值寿命季节性表现为:秋季夏季春季,并且在各个季节均表现,高级根寿命显著大于一级根寿命(P0.01);一级根仅夏季与秋季差异性不显著(P0.05),而高级根仅春季与秋季存在极显著差异(P0.01);(3)一级根和高级根距树干基部0 cm处细根中值寿命均大于50 cm处一级根和高级根细根的中值寿命。同一位置处高级根寿命要大于一级根寿命。在距树干基部0 cm处和50 cm处,一级根和高级根的寿命均存在极显著差异(P0.01),但高级根却在距树干基部0 cm和50 cm处差异不明显,而一级根却表现极显著差异(P0.01)。     

塔克拉玛干沙漠腹地梭梭幼苗根系分布特征对不同灌溉量的响应

在塔克拉玛干沙漠腹地,采用分层分段挖掘法对不同灌溉量条件下(每株每次灌水35、24.5和14 kg)梭梭(Haloxylon ammodendron)幼苗根系的分布特征进行了研究。结果表明: 1)随着灌溉量的减少,梭梭幼苗根系生物量的分布格局有向深层发展的趋势,在不同灌溉量条件下地下垂直各层生物量与土壤垂直深度呈显著的负对数关系;2)各灌溉量梭梭幼苗的最大水平根长为垂直根长的2倍,但不同灌溉量根系生物量的水平分布趋势一致;3)吸收根生物量的垂直分布与土壤含水量的垂直变化基本一致,均呈“单峰型”曲线,但灌溉量不同,吸收根生物量峰值在土壤中出现的位置也不同,随着灌溉量的减少,吸收根集中分布区有向深层发展的趋势;4)根长、根表面积和根体积随着土壤深度的增加均呈“单峰型”曲线,灌溉量愈小,根长、根表面积和根体积的峰值愈位于土壤的深层;5)根冠比和垂直根深与株高之比随着灌溉量的减少而呈增加的趋势。     

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。     

Evaluation in field conditions of a three-dimensional architectural model of the maize root system: Comparison of simulated and observed horizontal root maps

Most existing water and nutrient uptake models are based on the assumption that roots are evenly distributed in the soil volume. This assumption is not realistic for field conditions, and significantly alters water or nutrient uptake calculations. Therefore, development of models of root system growth that account for the spatial distribution of roots is necessary.The objective of this work was to test a three dimensional architectural model of the maize root system by comparing simulated horizontal root maps with observed root maps obtained from the field. The model was built using the current knowledge on maize root system morphogenesis and parameters obtained under field conditions. Simulated root maps (0.45 × 0.75 m) of horizontal cross sections at 3 depths and 3 dates were obtained by using the model for a plant population. Actual root maps were obtained in a deep, barrier-free clay-loamy soil by digging pits, preparing selected horizontal planes and recording root contacts on plastic sheets.Results showed that both the number of cross-sections of axile roots, and their spatial distribution characterized with the R-index value of Clark and Evans (1954), were correctly accounted for by the model at all dates and depths. The number of cross-sections of laterals was also correctly predicted. However, laterals were more clustered around axile roots on simulated root maps than on observed root maps. Although slight discrepancies appeared between simulated and observed root maps in this respect, it was concluded that the model correctly accounted for the general colonization pattern of the soil volume by roots under a maize crop.     

Effect of soil mechanical impedance on root growth of two rice varieties under field drought stress

Two upland rice varieties, Azucena and Bala, were screened for root growth under droughted and irrigated treatments in two field sites at the West Africa Rice Development Association (WARDA) experimental farm, Côte d’Ivoire, during the dry season of 1999/2000. The sites were chosen to represent contrasting soil profile penetration resistance (PR) characteristics on upland sites, although both were relatively impeding. The number of nodal root axes per unit area passing through horizontal transects (root density) was counted at 35, 56, 77 and 98 days after sowing (DAS) at 10 cm depth intervals. Azucena consistently maintained a greater root density than Bala and a greater proportion of Azucena roots grew to 30 cm depth (22.7% vs. 8.4% at 77 DAS). There was little detectable effect of water regime on root distribution but evidence of lower root numbers at depths below 20 cm in the higher PR site was revealed. A site by variety by soil depth interaction suggests that Azucena roots are more strongly affected by very high PR than those of Bala. PR between 0–30 cm depth increased greatly with decreasing soil water content during the drought as the soil dried. This increase is likely to have prevented or greatly impaired further nodal root growth within this layer. At 40 cm depth, PR was high (3–4 MPa) but did not increase during the drought. At this depth root growth rate was likely to be greatly reduced despite the availability of water. These results demonstrate that varietal differences in root morphology characterised in the laboratory can be also detected in impeding field soils as differences in the density of roots at depth. Relatively poor root growth in these fields in the absence of drought was probably due to the high mechanical impedance and/or the physiological stress of the plants in the dry season. Our results indicate that high mechanical impedance was a more fundamental constraint on root growth than soil water availability during the drought. Thus, varietal differences in root penetration ability might be very important for drought avoidance in soils of this type.     

星毛委陵菜根系构型对草原退化的生态适应

对轻度、中度、重度和极度退化的草原群落中星毛委陵菜(Potentilla acaulis)根系构型参数及相应的土壤水分、容重和硬度等指标进行了分析, 以研究星毛委陵菜根系构型对草原退化的生态适应性。结果表明: 1)在以大针茅(Stipa grandis)为建群种的典型草原中, 随着退化程度的加剧, 星毛委陵菜在群落中的作用逐渐增强, 其根幅、根深、一级垂向根数、分蘖子株数和水平分蘖根长度显著增加; 2)根表面积、二级侧根长度、总根长和根分叉数4个根系构型参数是解释星毛委陵菜根系构型对草原退化生态适应的首选指标, 解释力依次减小, 累计贡献率为92.34%; 3)直径2 mm以下的根系对单株系星毛委陵菜的根表面积和总根长影响显著; 4)阔腰倒锥体三维根系构型是星毛委陵菜适应草原退化并使之成为建群种的优势构型。     

Number,position, diameter and initial direction of growth of primary roots in Musa

To understand soil colonization by a root system, information is needed on the architecture of the root system. In monocotyledons, soil exploration is mainly due to the growth of adventitious primary roots. Primary root emergence in banana was quantified in relation to shoot and corm development. Root emergence kinetics were closely related to the development of aerial organs. Root position at emergence on the corm followed an asymptotic function of corm dry weight, so that the age of each root at a given time could be deduced from its position. Root diameter at emergence was related to the position of the roots on the corm, with younger roots being thicker than older ones. However, root diameters were not constant along a given root, but instead decreased with the distance to the base; roots appear to be conical in their basal and apical parts. Root growth directions at emergence were variable, but a high proportion of the primary roots emerged with a low angle to the horizontal. Further research is needed to evaluate whether these initial trajectories are conserved during root development. Results presented in this study are in good agreement with those reported for other monocotyledons such as maize and rice. They give quantitative information that will facilitate the development of models of root system architecture in banana.     

三峡库区马尾松根系生物量的空间分布

以三峡库区主要植被马尾松人工林为研究对象,用内径为10 cm的根钻,分别在马尾松中龄林、近熟林和成熟林内,据树干0.5、1.0、1.5 m和2.0 m处设置取样点,各样点按0-10、10-20、20-30、30-40、40-60 cm将土壤分为5个垂直层次,对马尾松根系的空间分布格局进行调查。结果表明:(1)三峡库区马尾松总根系生物量(0-10 mm)为中龄林(4.72 t/hm²)显著高于成熟林(2.94 t/hm²)和近熟林(2.40 t/hm²)(P<0.05)。细根(0-2 mm)生物量随年龄增加而递减,差异不显著(P>0.05);(2)马尾松3个林龄中根系生物量表现出一定的水平分布特征,但具体趋势表现各异,细根生物量最大值均出现在距离样木1.0 m处;(3)细根主要分布在土壤上层,其中47.53%-71.73%的活细根集中在0-20 cm土壤深度内,且随土层的加深,其生物量明显减少。粗根(2-10 mm)则主要分布于20-60 cm土层范围内;(4)根系直径越小,受环境变化越明显。马尾松细根生物量分布主要受土壤深度的影响,树龄和不同水平距离对细根分布格局影响不显著(P>0.05),各因素对粗根生物量的影响均未达到显著水平(P>0.05)。     

种间互作和施氮对蚕豆/玉米间作生态系统地上部和地下部生长的影响

为河西走廊绿洲灌区豆科/禾本科间作体系的养分管理提供科学依据,于2007年在武威绿洲农业试验站应用田间原位根系行分隔技术研究了蚕豆/玉米种间互作和施氮对玉米抽雄期的根系空间分布、根系形态和作物地上部生长的影响。研究结果表明:种间互作和施氮均增加了玉米和蚕豆在纵向和横向两个尺度上的根重密度、根长密度、根表面积、根系体积。根长密度和根表面积与两种作物产量和氮素吸收均呈正相关,而与蚕豆的根瘤重呈负相关;抽雄期的土壤含水量与玉米产量和养分吸收呈显著的负相关。玉米根系可以占据蚕豆地下部空间,但蚕豆的根却较少到间作玉米的地下部空间,也就是间作后增加了玉米根系水平尺度的生态位。蚕豆和玉米根系主要分布分别在0-40 cm浅土层和0-60 cm 土层,且间作玉米根系在60-120 cm比单作和分隔的多。因此,种间互作和施氮扩大了两作物根系纵向和横向的空间生态位,改变了作物根系形态,即扩展了两者水分和养分吸收的生态位,增加了作物吸收养分的有效空间,从而提高了间作生态系统的生产力。     

Vertical distribution of maize roots in relation to permanent soil characteristics

The vertical distribution of maize roots was studied in four contrasting soils, (arenosols, luvisols, planosols and vertisols) by using in-situ root mapping on vertical planes. The relationship between root contact frequency and depth was different for each soil, with a relatively low field-to-field variability within each soil type. The general aspect of this relationship did not change appreciably for three years in arenosols, with a low colonization in sandy layers probably being due to mechanical barriers. The relationship was consistently non-monotonic in luvisols and planosols, because of the sparse colonization of sandy layers. In planosols, these layers were traversed by some primary roots, which were essentially clustered in animal burrows. The distribution of root contact frequency was closer to an exponential function in vertisols. In these soils rooting depth and colonization of deep soil layers showed a marked increase during two dry years compared with a wet year. This was probably due to a denser net of shrinkage cracks and slickensides, where roots were essentially located in dry years. These results raise the possibility of modelling the decrease in root distribution with depth using soil information and climatic characteristics.     

Stand fine root biomass and fine root morphology in old-growth beech forests as a function of precipitation and soil fertility

Only very limited information exists on the plasticity in size and structure of fine root systems, and fine root morphology of mature trees as a function of environmental variation. Six northwest German old-growth beech forests (Fagus sylvatica L.) differing in precipitation (520 – 1030 mm year^–1) and soil acidity/fertility (acidic infertile to basic fertile) were studied by soil coring for stand totals of fine root biomass (0–40 cm plus organic horizons), vertical and horizontal root distribution patterns, the fine root necromass/biomass ratio, and fine root morphology (root specific surface area, root tip frequency, and degree of mycorrhizal infection). Stand total of fine root biomass, and vertical and horizontal fine root distribution patterns were similar in beech stands on acidic infertile and basic fertile soils. In five of six stands, stand fine root biomass ranged between 320 and 470 g m^–2; fine root density showed an exponential decrease with soil depth in all profiles irrespective of soil type. An exceptionally small stand fine root biomass (<150 g m^–2) was found in the driest stand with 520 mm year^–1 of rainfall. In all stands, fine root morphological parameters changed markedly from the topsoil to the lower profile; differences in fine root morphology among the six stands, however, were remarkably small. Two parameters, the necromass/biomass ratio and fine root tip density (tips per soil volume), however, were both much higher in acidic than basic soils. We conclude that variation in soil acidity and fertility only weakly influences fine root system size and morphology of F. sylvatica, but affects root system structure and, probably, fine root mortality. It is hypothesized that high root tip densities in acidic infertile soils compensate for low nutrient supply rates, and large necromasses are a consequence of adverse soil chemical conditions. Data from a literature survey support the view that rainfall is another major environmental factor that influences the stand fine root biomass of F. sylvatica.     

柠条细根的空间分布特征及其季节动态

以晋西北黄土区30年生柠条(Caragana korshinskii Kom.)人工林为研究对象,2007年应用Minirhizotron技术,分别在距茎干水平距离0、50、100 cm处设点,对林地0-100 cm土层深度范围内的柠条细根空间分布及其生长季的动态进行了研究。结果表明:(1)生长季柠条细根根长密度(RLD)总平均值为1.3423 mm/cm²。在水平方向上,距茎干水平距离50 cm处分布最多(1.5369 mm/cm²),其次为0 cm处(1.3855 mm/cm²), 100cm处分布最少(1.1044 mm/cm²)。在垂直深度上,各土层RLD平均值大小顺序为40-60 cm>60-80 cm>20-40 cm>0-20 cm>80-100 cm;(2)在0-100 cm土层范围内,月平均RLD在生长季的波动范围为0.4405 2.1040 mm/cm²,其中9月份最多,4月份最少;RLD在5个土层深度3个水平距离处随季节变化均表现先增加后减少的趋势,且不同空间位置RLD峰值变化均在秋季(8 10月份)波动。细根的这种时空分布差异,可能主要受林下土壤资源空间异质性及其季节性变化的影响,但也不排除其它因素的影响(如真菌,植食性昆虫)。     

塔里木河下游阿拉干断面胡杨根系空间分布规律研究

采用根系质量、密度、消弱系数和表面积等指标对塔里木河下游主要建群种胡杨根系空间分布特征进行定量研究,并分析了胡杨根系分布与土壤含水率的关系,为塔里木河下游受损天然胡杨林恢复提供依据。研究表明:(1)细根分布频数占活根总数的比例最大,死根所占比例从表层到深层有逐渐递减的趋势。(2)单株胡杨根系密度分布不对称。随着水平距离的增加,细根根系长度呈现近长远短的特点,而根量的水平分布呈现近重远轻的特点,主要集中在100～200 cm范围内。(3)胡杨根系垂直分层明显,根量在垂直方向上呈两头小、中间大的梭形分布,根系消弱系数值较大。(4)在土层30～120 cm范围内,细根表面积分布与土壤含水率分布呈显著正相关。     

A comparison of 14C, 86Rb,and total excavation for determination of root distributions of individual plants

Individual grass (Bouteloua gracilis) and shrub (Gutierrezia sarothrae) plants were either excavated as monoliths on nail boards, exposed to ¹⁴CO₂, or stem-injected with ⁸⁶Rb to compare the ability of the techniques to determine horizontal and vertical distribution of roots. The vertical distribution or roots directly under plant centers obtained by coring most closely correlated with monolith root length. ¹⁴C activity greatly overestimated near-surface roots and underestimated deep roots. ⁸⁶Rb activity did not follow the pattern of geometric decrease in root biomass with depth. Comparisons of both isotopes with monolith root length, over both horizontal and vertical axes, indicated that ¹⁴C activity was consistently concentrated near the soil surface, and ⁸⁶Rb activity was highly variable and randomly distributed. ¹⁴C may better represent root activity than root mass, and stem-injection methods can result in nonuniform labeling of roots. Caution should be exercised when using tracers to infer root biomass distributions. Resource partitioning between shrubs and grasses is discussed in relation to soil water dynamics in this semiarid grassland.     

典型黑土区主要树种根系构型特征及其对固土能力的影响

为量化典型黑土区主要树种根系构型特征,探究其对固土能力的影响,以该区分布较广的榆叶梅、小叶锦鸡儿、白桦、糖槭、红皮云杉、樟子松单株个体为研究对象,采用全根挖掘和WinRHIZO Pro LA2004分析系统相结合对其根系空间分布、几何形态、分形等特征进行测定,同时采用原位整株根系拉拔的方法量化根系垂直拉拔力。结果表明: 榆叶梅以倾斜根为主,小叶锦鸡儿、白桦、糖槭和红皮云杉以水平根为主,樟子松根系在水平和垂直分布上较为均衡;除白桦总根表面积和红皮云杉总根长外,灌木树种总根长、总根表面积显著大于乔木,落叶阔叶乔木总根长、总根表面积显著大于针叶常绿乔木,白桦总根体积显著大于小叶锦鸡儿、糖槭、红皮云杉和樟子松;榆叶梅、小叶锦鸡儿和白桦根系分形维数和分形丰度显著大于红皮云杉和樟子松;榆叶梅、小叶锦鸡儿和糖槭整株根系平均最大垂直拉拔力显著大于白桦、樟子松和红皮云杉。主要受根系总根长、总根表面积和倾斜根数量的影响,榆叶梅、小叶锦鸡儿和糖槭根系表现出较强的固土能力,可作为典型黑土区水土保持植被构建中优先选择的树种。     

Fine Root Distribution in Dehesas of Central-Western Spain

A Dehesa is a structurally complex agro-silvo-pastoral system where at least two strata of vegetation, trees and herbaceous plants coexist. We studied the root distribution of trees (Quercus ilex L.) and herbaceous plants, in order to evaluate tree and crops competition and complementarity in Dehesas of Central Western Spain. 72 soil cores of 10 cm diameter (one to two metre deep) were taken out around 13 trees. Seven trees were intercropped with Avena sativa L. and six trees were in a grazed pasture dominated by native grasses. Soil coring was performed at four distances from the tree trunks, from 2.5 (beneath canopy) till 20 m (out of the canopy). Root length density (RLD) of herbaceous plants and trees was measured using the soil core-break method. Additionally, we mapped tree roots in 51 profiles of 7 recently opened road cuts, located between 4 and 26 m of distance from the nearest tree. The depth of the road cuts varied between 2.5 and 5.5 m. Herbaceous plant roots were located mostly in the upper 30 cm, above a clayey, dense soil layer. RLD of herbaceous plants decreased exponentially with depth until 100 cm depth. Holm-oak showed a much lower RLD than herbs (on average, 2.4 versus 23.7 km m⁻³, respectively, in the first 10 cm of the soil depth). Tree RLD was surprisingly almost uniform with depth and distance to trees. We estimated a 5.2 m maximum depth and a 33 m maximum horizontal extension for tree roots. The huge surface of soil explored by tree roots (even 7 times the projection of the canopy) could allow trees to meet their water needs during the dry Mediterranean summers. The limited vertical overlap of the two root profiles suggests that competition for soil resources between trees and the herbaceous understorey in the Dehesa is probably not as strong as usually assumed.     

Root distribution and morphology of maize seedlings as affected by tillage and fertilizer placement

Suboptimal soil conditions are known to result in poor early growth of maize (Zea mays L.) in no-tillage (NT) systems in contrast with conventional tillage (CT) systems. However, most studies have generally focused on maize roots at later growth stages and/or do not give details on root morphology. In a 2-year field study at two locations (silt loam and loam soils) in the Swiss midlands, we investigated the impacts of tillage intensity, NT vs. CT, and NP-fertilizer sidebanding on the morphology, vertical and horizontal distribution, and nutrient uptake of maize roots at the V6 growth stage. The length density (RLD) and the length per diameter-class distribution (LDD) of the roots were determined from soil cores taken to a depth of 0.5 m and at distances of 0.05 and 0.15 m from both sides of the maize row. The temperature of the topsoil was lower, and the bulk density and penetration resistance were greater in the topsoil of NT compared with CT. The growth and the development of the shoot were slower in NT. RLD was greater and the mean root diameter smaller in CT than in NT, while the vertical and horizontal distribution of roots did not differ between CT and NT. RLD increased in the zone enriched by the sidebanded fertilizer, independent of the tillage system, but LDD did not change. The poorer growth of the roots and shoots of maize seedlings was presumably caused by the lower topsoil temperature in NT rather than by mechanical impedance. The placement of a starter fertilizer at planting under NT is emphasized.     

黄河三角洲滨海盐碱地11个造林树种细根解剖性状对土壤条件的响应

细根作为植物与土壤连接的重要部位,能够反映植物对生存环境的适应性。以黄河三角洲滨海盐碱地不同立地条件下11个造林树种为对象,基于细根分支等级划分1-4级根序并进行解剖特征测定,分析细根解剖性状对滨海盐碱地不同土壤条件的响应规律。结果表明：（1）不同根序的细根直径存在显著差异,细根直径随根序升高呈增大趋势,而同根序的细根直径在不同树种间表现出显著的种间差异（P < 0.05）。1-2级细根皮层厚度、3-4级细根导管密度在树种间的差异均达显著水平（P < 0.05）。（2）在较为严重盐渍化土壤条件下（立地1）,细根皮层厚度较其他立地显著增大,但细根导管密度较小;在轻度盐碱立地条件下（立地3）,细根导管密度较大;较为严重的盐碱立地具有更为发达的细根直径及维管柱直径。（3）树种1-2级细根解剖结构与土壤环境关系最为密切,其中1级根直径与土壤pH值显著正相关（P < 0.05）,与土壤硝态氮含量呈显著负相关（P < 0.05）。对土壤理化性质与细根解剖性状的冗余分析表明,前两个轴的特征值达0.640和0.196,土壤速效养分含量与轴一（RDA1）呈正相关,低级根解剖性状则与轴二（RDA2）呈显著负相关。低级根解剖结构以及土壤的pH值能解释较多树种的差异性,其中低级根直径与皮层厚度对盐碱环境表现出较强的响应。     

Effect of soil water content on the gravitropic behavior of nodal roots in maize

The direction of root growth is an important factor that determines the spatial distribution of roots in the soil. The influence of soil water content on the direction of growth of maize nodal roots was studied both in the field and in the greenhouse. In the field experiment, the one plot was regularly irrigated (I-plot) and the other non-irrigated (N-plot). In the greenhouse experiment, three water treatments were conducted on plants grown in pots: continuously wet (CW), early drying (ED), and late drying (LD). The direction of root growth was quantified by the angle from the vertical, measured at 1 cm intervals for 10 cm from the first five internodes. Nodal roots grew more vertically in the N-plot and ED treatment than those in the I-plot and CW treatment. This was due to the decrease of the initial angle and/or the liminal angle. It is therefore thought that two events regulate the growth direction of nodal roots under dry soil conditions: gravitropic bending at root emergence from the stem and the later establishment of the angle of growth. Nodal roots appearing after rewatering in the ED treatment grew in a similar direction as those in the CW treatment. It follows from this that the water content of the surrounding soil has a direct effect on the direction of growth. Nodal roots that emerged in rapidly drying soil in the LD treatment ceased growing after showing negative gravitropism. The possible mechanisms determining the growth direction of nodal roots in drier soils are discussed.