Mapping QTLs associated with drought avoidance in upland rice grown in the Philippines and West Africa

Localizing genes that contribute to drought avoidance in a quantitative way should enable the exploitation of these genes in breeding through marker-assisted selection, and may lead to the discovery of gene identity and function. Between 110 and 176 F₆ recombinant inbred lines from a mapping population derived from a cross of upland rice varieties Bala and Azucena have been evaluated for indicators of drought avoidance in sites in the Philippines and West Africa over two dry seasons. A molecular map with 102 RFLP, 34 AFLP and six microsatellite markers has been used to map (by composite interval mapping) quantitative trait loci (QTLs) for the visual scores of leaf rolling and leaf drying and leaf relative water content. QTLs were mapped for each site and across sites. A total of 17 regions were identified which contained QTLs with a LOD score greater than 3.2. For leaf rolling, Bala was the parent contributing the majority of positive alleles whilst for the other traits, Bala and Azucena contributed more evenly. Six of the 17 regions influenced more than one trait, explaining the phenotypic correlations between traits that were observed. Three QTLs appeared to be specific to the Philippines experiments. One QTL had opposing effects in the Philippines and West Africa. QTLs for relative water content were detected on chromosome 8, congruent with an osmotic adjustment QTL identified in another population. Only three of the QTLs identified here have not been reliably identified in the two other populations that have been screened for drought avoidance. By using several populations assessed for drought avoidance in different sites, the distribution and utility of QTLs for drought avoidance in rice is being elucidated.     

Mapping QTLs and candidate genes for rice root traits under different water-supply conditions and comparative analysis across three populations

To investigate the genetic factors underlying constitutive and adaptive morphological traits of roots under different water-supply conditions, a recombinant inbred line (RIL) population derived from a cross between the lowland rice variety IR1552 and the upland rice variety Azucena with 249 molecular markers, was used in cylindrical-pot experiments. Eighteen QTLs were detected for seminal root length (SRL), adventitious root number (ARN), and lateral root length (LRL) and lateral root number (LRN) on the seminal root at a soil depth of from 3 to 6 cm under flooding and upland conditions. One identical QTL was detected under both flooding and upland conditions. The relative parameters under the two water-supply conditions were also used for QTL analysis. Five QTLs for upland induced variations in the traits were detected with the positive alleles from Azucena. A comparative analysis was performed for the QTLs detected in this study and those reported from two other populations with Azucena as a parent. Several identical QTLs for root elongation were found across the three populations with positive alleles from Azucena. Candidate genes were screened from ESTs and cDNA-AFLP clones for comparative mapping with the detected QTLs. Two genes for cell expansion, OsEXP2 and endo-1,4--D-glucanase EGase, and four cDNA-AFLP clones from root tissues of Azucena, were mapped on the intervals carrying the QTLs for SRL and LRL under upland conditions, respectively.Communicated by H.C. Becker     

Genotypic differences in root penetration ability of wheat through thin wax layers in contrasting water regimes and in the field

A significant proportion of arable land in south-western Australia is highly susceptible to subsoil compaction, which limits access of roots of wheat to water and nutrients at depth. Genotypic variation in the ability of roots to penetrate a hardpan has been reported for other cereals, using a pot technique, where a thin wax-layer of paraffin wax and petroleum jelly is placed in a soil column to simulate a hardpan. Previously we have modified and validated this technique for measuring root penetration ability of wheat seedlings under contrasting water regimes. Here we report on a series of five experiments (runs), two in well-watered and three in drought stress conditions, which evaluated seminal and nodal root penetration ability through thin wax layers among 24 Australian wheat cultivars and breeding lines (entries). These results were compared with observations on their rooting depths in two contrasting soil types in field trials, including a sandy duplex that contained a hardpan and a red clay that increased in soil strength with depth. Nodal roots ceased growth early under soil water deficit, and water uptake was instead dependant on seminal roots under conditions imposed in the pots. Plants were then reliant on the ability of seminal roots to penetrate the wax layer. Eight entries had superior root penetration ability in both well-watered and drought stressed conditions. Roots of three other entries, which failed to penetrate the wax layers, died under drought stress conditions. In field trials, there was a significant interaction between site and entry for maximum root depth. Our results from the pot studies and field trials indicate that there exists genotypic variation in root traits that are required to penetrate uniformly hard soil, dry soil or soil containing a hardpan. As four of the eight superior entries also showed superior root penetration ability at both sites in the field, there was an overall consistency, but there were exceptions at individual field sites. Factors likely to result in such exceptions were discussed, and topics for further research identified.     

Assessing the importance of genotype x environment interaction for root traits in rice using a mapping population. I: a soil-filled box screen

Altering root system architecture is considered a method of improving crop water and soil nutrient capture. The analysis of quantitative trait loci (QTLs) for root traits has revealed inconsistency in the same population evaluated in different environments. It must be clarified if this is due to genotype × environment interaction or considerations of statistics if the value of QTLs for marker-assisted breeding is to be estimated. A modified split-plot design was used where a main plot corresponded to a separate experiment. The main plot factor had four treatments (environments), which were completely randomized among eight trials, so that each treatment was replicated twice. The sub-plot factor consisted of 168 recombinant inbreed lines of the Bala × Azucena rice mapping population, randomly allocated to the seven soil-filled boxes. The aim of the trial was to quantify QTL × environment interaction. The treatments were chosen to alter partitioning to roots; consisting of a control treatment (high-soil nitrogen, high light and high-water content) and further treatments where light, soil nitrogen or soil water was reduced singly. After 4 weeks growth, maximum root length (MRL), maximum root thickness, root mass below 50 cm, total plant dry mass (%), root mass and shoot length were measured. The treatments affected plant growth as predicted; low nitrogen and drought increased relative root partitioning, low-light decreased it. The parental varieties Bala and Azucena differed significantly for all traits. Broad-sense heritability of most traits was high (57–86%). Variation due to treatment was the most important influence on the variance, while genotype was next. Genotype × environment interaction was detected for all traits except MRL, although the proportion of variation due to this interaction was generally small. It is concluded that genotype × environment interaction is present but less important than genotypic variation. A companion paper presents QTL × environment analysis of data.     

根区交替控制灌溉条件下玉米根系吸水规律

为了阐明根区交替控制灌溉(CRDAI)条件下玉米根系吸水规律,通过田间试验,在沟灌垄植模式下采用根区交替控制灌溉研究玉米根区不同点位(沟位、坡位和垄位)的根长密度(RLD)及根系吸水动态。研究表明,根区土壤水分的干湿交替引起玉米RLD的空间动态变化,在垄位两侧不对称分布,并存在层间差异;土壤水分和RLD是根区交替控制灌溉下根系吸水速率的主要限制因素。在同一土层,根系吸水贡献率以垄位最大,沟位最低;玉米营养生长阶段,10—30 cm土层的根系吸水速率最大;玉米生殖生长阶段,20—70 cm为根系吸水速率最大的土层,根系吸水贡献率为43.21%—55.48%。研究阐明了交替控制灌溉下根系吸水与土壤水分、RLD间相互作用的动态规律,对控制灌溉下水分调控机理研究具有理论意义。     

A combined RFLP and AFLP linkage map of upland rice (Oryza sativa L.) used to identify QTLs for root-penetration ability

Acombined RFLP and AFLP linkage map of an F₆ recombinant inbred population, which was derived from a previously mapped F₂ of a cross between the two drought resistant upland rice varieties Bala and Azucena, is presented. The map contains 101 RFLP and 34 AFLP markers on 17 linkage groups covering 1680 cM. Also presented is the approximate mapping position of a further four RFLP and 75 AFLP markers, which either could not be given a unique place on the map or for which the available data is not sufficient to allow confident positioning, and the result of quantitative trait locus (QTL) mapping of traits related to root-penetration ability. Root penetration was assessed by counting the number of root axes that penetrated a 3 mm-thick layer consisting of 80% wax and 20% white soft paraffin. Good root penetration would be expected to increase drought resistance where soil strength is high. Single-marker analysis revealed seven QTLs for the number of roots which penetrate the wax layer. In identical locations were seven QTLs for the ratio of penetrated to the total number of roots. Transgressive inheritance of positive alleles from Bala explained four of these QTLs. Comparison of the QTLs identified here with previous reports of QTLs for root morphology suggest that alleles which improve root penetration ability may also either make the roots longer or thicker. Received: 3 February 1999 / Accepted: 30 April 1999     

Yield response to water deficit in an upland rice mapping population: associations among traits and genetic markers

A population of recombinant inbred rice lines from a cross between the upland japonica cultivar Azucena and the upland indica cultivar Bala was evaluated in a series of upland field experiments. Water stress was imposed during the reproductive stage by managed irrigation during the dry season, while control treatments were maintained in aerobic, well-irrigated conditions. Water deficit resulted in a yield reduction of 17 to 50%. The genetic correlation between stress and control yields was quite high when stress was mild, and the heritability of yield was similar in stress and control treatments across both years of this study. Genetic correlations between secondary traits such as leaf rolling and drying and yield under stress varied from high (leaf drying) to insignificant (leaf rolling). Lines with superior yield tended to have fewer panicles and larger grain size than the high-yielding parent, Bala, even though the panicle number was positively correlated with yield and the thousand-grain weight was not associated with yield for the population as a whole. Analysis of quantitative trait loci (QTLs) for yield and yield components allowed the identification of 31 regions associated with growth or yield components. Superior alleles came from either parent. Several of the regions identified had also been reported for root mass at depth or maximum root length in this population in other studies made under controlled environments, and for leaf drying (LD) in field studies. However, the direction of the effect of QTLs was not consistent, which indicates that there was not necessarily a causal relationship between these secondary traits and performance. We conclude that mapping populations can provide novel insights on the actual relationships between yield components and secondary traits in stress and control environments and can allow identification of significant QTLs for yield components under drought stress.Abbreviations DAS Days after sowing - GPP Grains per panicle - QTL Quantitative trait locus - RWC Relative water content - SPP Spikelets per panicle - TGW Thousand-grain weight - VPD Vapor pressure deficit     

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.     

土壤水分梯度对白三叶(Trifolium repens)光合作用和根系分布的影响

通过对两个品种白三叶Trifolium repens cv.Haifa(海发)和Trifolium repens cv.Rivendel(瑞文德)盆栽试验,模拟3种不同的土壤水分状况(无水分胁迫:保持植株良好的水分供应;轻度胁迫:表层0～20cm土壤处于干旱状态;重度胁迫:表层0～20cm土壤处于极干旱状态,20～40cm土壤处于干旱状态)对白三叶光合作用和根系生长的影响.结果表明,当植株未遭受水分胁迫时,两个品种白三叶的光合作用和根系生长状况没有明显差异;当表层0～20cm处于干旱状态时,'海发'在处理后期的净光合速率和水分利用效率升高,根系生长量增大,表现出促进作用,'瑞文德'受到的影响不显著;当表层0～20cm处于极干旱、20～40cm处于干旱状态时,'海发'在处理前期受到轻微影响,随后恢复正常状态,'瑞文德'则受到较严重的影响.随着干旱程度的加深和时间的延长,白三叶的根冠比逐渐增大.与'瑞文德'相比,在相同时期相同胁迫程度下,'海发'的根冠比没有显著差异,但深根数量大大超过'瑞文德',因而,'海发'的耐旱能力强于'瑞文德'.     

乌兰布和沙区紫花苜蓿根系生长及吸水规律的研究

研究了不同水分处理下,乌兰布和沙区紫花苜蓿系生长发育规律及根系吸水速度,结果表明：不同水分处理的根系生长规律最基本一致的,在生长季风均呈增加的趋势;但适度干旱可促进根系的伸长生长。在当地土壤类型条件下,根系主要分布在０－３０cm土层;根重密度在土壤剖面上的分布遵循对数规律,并随深度的增加呈降低趋势。运用一维土壤水分运动方程,计算得到了不同水分处理根系吸水速度在土壤剖面上的分布状况;根系吸水速率与土壤含水量和根密度密切相关。     

Root growth and nitrate uptake by wheat (Triticum aestivum L.) following wetting of dry surface soil

The nitrate uptake capacity of surface roots of spring wheat(Triticum aestivum L. cv. Kulin) was investigated followingwetting of dry surface soil. Plants were grown to stem elongationstage with adequate watering at depth while the surface soilwas allowed to dry. Eight weeks after sowing, water or a ¹⁵N-nitratesolution was added to the surface soil to simulate rainfall.Root growth and nitrate uptake were measured up to 4 d afterwetting on plants with unconfined nodal root growth and on plantswith the majority of nodal roots confined within small vials.Prior to wetting, plants from both nodal treatments had seminalroots with collapsed cortices along the upper 10 cm and manyshort, viable lateral roots. Nodal roots, where present, wereonly a few cm long and unbranched. Only plants with unconfinednodal roots were able to take up nitrate within the 24 h beforeany new root growth. By 2 d after wetting there was significantgrowth of the seminal lateral roots, and rapid growth and branchingof nodal roots. From 2 d after wetting, plants with confinednodal roots also took up nitrate, presumably due to the growthof the seminal lateral roots. Hence it appears as though thenodal roots in the unconfined treatment could immediately takeup nitrate, but the seminal roots required new lateral rootgrowth to become active in nitrate uptake. The plants with confinednodal roots had a lower nitrate uptake than those with unconfinednodal roots 4 d after wetting, indicating that the seminal rootsystem was not able to compensate for lack of nodal roots. Insufficientnitrate was taken up after 4 d, by plants from either nodalroot treatment, to increase the shoot N concentration significantly. Key words: Triticum aestivum, nitrate uptake, drought, seminal roots, nodal roots     

不同抗旱性花生品种的根系形态发育及其对干旱胁迫的响应

为明确不同抗旱性花生品种的根系形态发育特征,探讨其根系形态发育特征对不同土壤水分状况的响应机制,在防雨棚旱池内进行土柱栽培试验,研究抗旱型品种“花育22号”、“唐科8号”和干旱敏感型品种“花育23号”3个不同抗旱性花生品种根系形态发育特征及其对干旱胁迫的响应.结果表明:抗旱型品种根系较发达,具有较大的根系生物量、总根长、总根系表面积.干旱胁迫使抗旱型品种根系总表面积和体积增加,而干旱敏感型品种则相反.干旱胁迫显著增加抗旱型品种“花育22号”20 cm以下土层内根长密度分布比例及根系表面积和体积,但“唐科8号”相应根系性状仅在20-40 cm土层内增加;干旱胁迫使干旱敏感型品种“花育23号”40 cm以下土层内各根系性状升高,但未达显著水平且其深层土壤内各根系性状增加幅度小于“花育22号”.花生根系总长、总表面积及0-20 cm土层内根系性状与产量间呈显著或极显著正相关.土壤水分亏缺条件下,花生主要通过增加深层土壤内根长、根系表面积和体积等形态特性,优化空间分布构型,以调节植株对水分的利用.     

A gradual rather than abrupt increase in soil strength gives better root penetration of strong layers

The purpose of this paper is to test the hypothesis that gradual rather than abrupt increases in strength lead to increased root penetration of strong layers. This was tested experimentally in a model system with multi-layered wax discs with a total thickness of up to 6 mm that either increased in strength with depth, or decreased in strength with depth. Strength was varied by altering the proportions of hard paraffin wax and white soft paraffin. Multi-layered wax discs consisted of three layers, each 1.5 or 2 mm thick, to give a disc thickness of either 4.5 or 6 mm. ‘Test’ wax discs had a 40% hard wax layer above a 60% wax layer, which was above an 80% wax layer. This led to a gradual increase in strength as roots encountered the disc from the sand growing medium. In ‘control’ discs, the order was reversed so that 80% wax was encountered immediately below the sand. These treatments were used to challenge roots of the rice cvs. Azucena and Bala. Compared with the control, the test layers increased root penetration of Azucena from a mean of 5.9 root axes to 12.4, while penetration of Bala was increased from 0.5 to 5.6. These results show that there was increased root penetration when roots encountered a gradual increase in strength rather than a sudden increase in strength. This was explored further with numerical simulations of stress distributions at the tips of roots. These simulations indicated that gradual rather than abrupt increases in strength decreased stress concentration at the root tip, consistent with the experimental results.     

Water relations and root activities of Buchloe dactyloides and Zoysia japonica in response to localized soil drying

Effects of localized soil drought stress on water relations, root growth, and nutrient uptake were examined in drought tolerant ‘Prairie’ buffalograss [Buchloe dactyloides (Nutt.) Engelm.] and sensitive ‘Meyer’ zoysiagrass (Zoysia japonica Steud.). Grasses were grown in small rhizotrons in a greenhouse and subjected to three soil moisture regimes: (1) watering the entire 80-cm soil profile (well-watered control); (2) drying 0–40 cm soil and watering the lower 40 cm (partially dried); (3) and drying the entire soil profile (fully dried). Drying the 0–40 cm soil for 28 days had no effect on leaf water potential (Ψ _leaf ) in Prairie buffalograss compared to the well-watered control but reduced that in Meyer zoysiagrass. Root elongation rate was greater for Prairie buffalograss than Meyer zoysiagrass under well-watered or fully dried conditions. Rooting depth increased with surface soil drying; with Prairie buffalograss having a larger proportion of roots in the lower 40 cm than Meyer zoysiagrass. The higher rates of water uptake in the deeper soil profile in the partially dried compared to the well-watered treatment and by Prairie buffalograss compared to Meyer zoysiagrass could be due to differences in root distribution. Root ¹⁵N uptake for Prairie buffalograss was higher in 0–20 cm drying soil in the partially dried treatment than in the fully dried treatment. Diurnal fluctuations in soil water content in the upper 20 cm of soil when the lower 40 cm were well-watered indicated water efflux from the deeper roots to the drying surface soil. This could help sustain root growth, maintain nutrient uptake in the upper drying soil layer, and prolong turfgrass growth under localized drying conditions, especially for the deep-rooted Prairie buffalograss. This revised version was published online in June 2006 with corrections to the Cover Date.     

Genotypic Variation in Root Growth Angle in Rice (Oryza sativa L.) and its Association with Deep Root Development in Upland Fields with Different Water Regimes

Deep root development, which is important for the drought resistance in rice (Oryza sativa L.), is a complex trait combining various root morphologies. The objective of this study was to elucidate genotypic variation in deep root development in relation to morphological indicators such as vertical root distribution and root growth angle. Two experiments were conducted: one on upland fields, and one in pots and fields. In experiment 1, the root systems of six rice cultivars on upland fields were physio-morphologically analyzed under different water regimes (irrigated and intermittent drought conditions during panicle development). In experiment 2, cultivar differences in root growth angles were evaluated with 12 cultivars using the basket method under irrigated conditions. No cultivar × environment interactions were found for total root length or deep root length between irrigated and drought conditions in experiment 1. This suggests that constitutive root growth, which is genetically determined, is important for deep root development under intermittent drought conditions during reproductive stage. Among root traits, the deep root ratio (i.e., deep root weight divided by total root weight) was most closely related to deep root length under both water regimes. This suggested that vertical root distribution constitutively affects deep root length. Significant genotypic variation existed in the nodal root diameter and root growth angle of upland rice in experiment 2. It was considered that genotypes with thick roots allocated more assimilates to deep roots through root growth angles higher to the horizontal plane on upland fields. This is the first report on genotypic variation in the root growth angle of rice on upland fields. It should prove useful for rough estimations of genotypic variation in the vertical root distribution of upland rice because root growth angle is rapidly and easily measured.     

Drought resistance and root morphology in sorghum

Two 2 m³ plots of soil were prepared to different water contents and each isolated from surrounding soil by impermeable plastic material. Nine sorghum varieties were germinated in the plots and allowed to grow without further watering. Time-to-wilt after emergence was measured, and several parameters relating to water flow of the seedling and nodal roots were determined. There was a good positive correlation between both seminal root and nodal root relative conductvity and time-to-wilt. In a second experiment, plants were germinated and grown in pots, and after two weeks of growth without further watering were inspected for survival in the unwilted state. The per cent survival was calculated. There was a negative correlation of seminal root relative conductivity with per cent survival, and a high negative correlation of the number of seminal roots with per cent survival. It is concluded that high relative conductivity indicates drought resistance if the plants are growing with less restricted roots as in open soil, while if the plants are grown in pots the reverse is the case. Experiments linking root conductivity with survival conducted in pots are poor predictors of performance in less restricted rooting conditions.     

干旱胁迫对花生根系生长发育和生理特性的影响

以花育17号和唐科8号两个花生品种为试验材料,在防雨棚栽培池内进行土柱栽培试验,研究了中度干旱胁迫和正常供水处理下花生生育后期根系形态发育特征和生理特性.结果表明: 唐科8号具有较发达的根系及较高的产量和抗旱系数,花育17号根系对干旱胁迫的适应性小于唐科8号.两品种根长密度、根系生物量均主要分布于0～40 cm土层中,但同一土层内两品种根系性状存在差异.与正常供水处理相比,干旱胁迫处理使花育17号各生育期总根长、根系总表面积和总体积均降低,而唐科8号除花针期显著降低外,其余生育期均明显升高;干旱胁迫增加了两品种20～40 cm土层内根系生物量、根系表面积和体积,而降低了40 cm以下土层内各根系性状;干旱胁迫处理使两品种饱果期40 cm以下土层内根系活力降低,且花育17号降低幅度高于唐科8号.干旱胁迫下两品种生育后期根系发育和生理特性的差异表明其根系在干旱胁迫下对水分吸收和利用存在差异.     

The impact of limited soil moisture and waterlogging stress conditions on morphological and anatomical root traits in maize (Zea mays L.) hybrids of different drought tolerance

Effects of soil drought or waterlogging on the morphological traits of the root system and internal root anatomy were studied in maize hybrids of different drought tolerance. The investigations comprised quantitative and qualitative analyses of a developed plant root system through determining the number, length and dry matter of the particular components of the root system and some traits of the anatomical structure of the seminal root. Obtained results have demonstrated a relatively broad variation in the habit of the root system. This mainly refers, to the number, length and dry matter of lateral roots, developed by seminal root, seminal adventitious and nodal roots as well as to some anatomical properties of the stele, cortex and metaxylem elements. Plants grown under waterlogging or drought conditions showed a smaller number and less dry matter of lateral branching than plants grown in control conditions. The harmful effect of waterlogging conditions on the growth of roots was greater when compared with that of plants exposed to drought. In the measurements of the root morphological traits, the effect of soil drought on the internal root anatomical characteristic was weaker than the effect of soil waterlogging. The observed effects of both treatments were more distinct in a drought sensitive hybrid Pioneer D than in drought resistant Pioneer C one. The drought resistant hybrid Pioneer C distinguished by a more extensive rooting and by smaller alterations in the root morphology caused by the stress conditions than drought sensitive hybrid Pioneer D one. Also the differences between the resistant and the sensitive maize hybrids were apparent for examined root anatomical traits. Results confirm that the hybrid Pioneer D of a high drought susceptibility was found to be also more sensitive to periodieal soil water excess. A more efficient water use and a lower shoot to root (S:R) ratio were found to be major reasons for a higher stress resistance of the hybrid Pioneer C. The reasons for a different response of the examined hybrids to the conditions of drought or waterlogging may be a more economical water balance and more favourable relations between the shoot and root dimensions in the drought resistant genotype. The observed modifications of the internal root structure caused by water deficit in plant tissues may partly influence on water conductivity and transport within roots. The results suggest that the morphological and anatomical traits of the maize root system may be used in practice as direct or indirect selection criteria in maize breeding.     

Hydraulic properties of sphagnum peat moss and tuff (scoria) and their potential effects on water availability

The importance of macrostructure to root growth of ryegrass (L. perenne) seedlings sown on the soil surface was studied in two soils in which the macrostructure had resulted mainly from root growth and macro-faunal activity. Sets of paired soil cores were used, one of each pair undisturbed and the other ground and repacked to the field bulk density. Undisturbed and repacked soils were first compared at equal water potentials in the range −1.9 to −300 kPa. At equal water potential, the undisturbed soil always had the greater strength (penetration resistance), and root growth was always greater in the repacked soil with no macrostructure than it was in the soil with macrostructure intact. At equal high strength (low water potentials) it appeared that root growth was better when soils were structured. When strength was low (high water potentials), root growth was better in the unstructured soil. Soils were then compared during drying cycles over 21 days. The average rate at which roots grew to a depth of 60 mm, and also the final percentage of plants with a root reaching 60 mm depth, was greatest in repacked soils without macrostructure. The species of vegetation growing in the soil before the experiment affected root growth in undisturbed soil; growth was slower where annual grasses and white clover had grown compared with soil which had supported a perennial grass. It appears that relatively few roots locate and grow in the macrostructure. Other roots grow in the matrix, if it is soft enough to be deformed by roots. Roots in the matrix of a structured soil grow more slowly than roots in structureless soil of equal bulk density and water potential. The development of macrostructure in an otherwise structureless soil, of the type studied, is of no advantage to most roots. However, once a macrostructure has developed, the few roots locating suitable macropores are able to grow at low water potential when soil strength is high. The importance of macrostructure to establishing seedlings in the field lies in rapid penetration of at least a few roots to a depth that escapes surface drying during seasonal drought. ei]{gnB E}{fnClothier}