Root Growth and Water Uptake by Maize Plants in Drying Soil

Sharp, R. E and Da vies, W. J. 1985. Root growth and water uptakeby maize plants in drying soil.— J. exp. Bot. 36: 1441–1456. The influence of soil drying on maize (Zea mays L.) root distributionand use of soil water was examined using plants growing in thegreenhouse in soil columns. The roots of plants which were wateredwell throughout the 18 d experimental period penetrated thesoil profile to a depth of 60 cm while the greatest percentageof total root length was between 20–40 cm. High soil waterdepletion rates corresponded with these high root densities.Withholding water greatly restricted root proliferation in theupper part of the profile, but resulted in deeper penetrationand higher soil water depletion rates at depth, compared withthe well watered columns. The deep roots of the unwatered plantsexhibited very high soil water depletion rates per unit rootlength. Key words: Maize, roots, water deficit, soil water depletion     

Stable isotope analysis of water extraction from subsoil in upland rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) as affected by drought and soil compaction

Water extraction from subsoil in upland rice (Oryza sativa L.) was examined as related to topsoil desiccation and subsoil compaction. The water extraction was observed by measurements of heavy water concentrations in transpiring plants. The plants were grown in pots that were filled with sandy soil and vertically compartmented into two columns. Heavy water was applied to the subsoil. Plants exposed to mild topsoil desiccation (–120 kPa in water potential) eventually increased water extraction from the subsoil and maintained photosynthetic rate and stomatal conductance at the wet condition level. The rates of the plants subjected to severely droughted topsoil (–190 kPa) were significantly lowered due to less water uptake from the subsoil. Subsoil compaction at bulk densities of 1.45 and 1.50 Mg m^–3 inhibited increase of root length densities. Limited water extraction from the subsoil was insufficient to maintain plant productivity under drought conditions. Daily water uptake per unit of root length in the lower tube did not apparently increase even if water demand on the unit root length increased. When water to topsoil was completely withheld, water extraction from the subsoil gradually increased as the topsoil dried out. Plants that were watered and rewatered took up very little water from the subsoil. The extraction from the subsoil occurred only when water potential of the topsoil was below about –190 kPa.     

Root distribution at various distances from clove trees growing in Indonesia

Efficient fertilizer application requires an understanding of the distribution of roots and soil nutrients in the soil profile. Cultural practices for clove trees in Indonesia has resulted in phosphorus (P) fertilizer being applied at the canopy edge; however, in these high P fixing soils efficient P fertilizer application should occur with the highest root densities. The objective of this study, therefore, was to determine the rooting distribution at various distances from the tree and soil depths for clove (Eugenia aromatica OK; variety Zanzibar) trees growing on an Andosol soil at Modoinding, Indonesia. Root distributions were determined to a 100-cm soil depth using soil cores at 0.5, 1.0 and 1.5 times the canopy radius for five 10-year-old clove trees grown on either level terrain or 23% slopes. Clove root length and weight densities decreased with soil depth and distance from the tree base. Fine clove roots (1 mm dia) comprised 72% of the total root length and was three to five times higher underneath the canopy than that outside the canopy. Roots were concentrated in the upper soil horizons; however, up to 36% of the total root length was found at a depth of 50–100 cm. Clove roots for trees growing at the level landscape position had the highest root length densities. Intercropped species root length densities were higher than clove root length densities at 1.5 times the canopy radius whereas intercropped root weight densities were higher than that for clove roots at both 1.5 and 1 times the canopy radius. Results suggest that fertilizer applications should be placed closer to the tree trunk rather than at the canopy edge to maximize P uptake by clove roots.     

Root distributions in a Grevillea robusta-maize agroforestry system in semi-arid Kenya

Limited knowledge of root distributions in agroforestry systems has resulted in assumptions that various tree species are more suited to agroforestry than others, because they are presumed to have few superficial lateral roots. This assumption was tested for Grevillea robusta when grown with maize (Zea mays) in an agroforestry system in a semi-arid region of Kenya. At a site with a shallow soil, root lengths of both species between the soil surface and bedrock were quantified by soil coring, at intervals over four cropping seasons, in plots containing sole stands and mixtures of the trees and crop; the trees were 4–6 years old and they were severely pruned before the third season. Profiles of soil water content were measured using a neutron probe. Prior to pruning of the trees, recharge of soil water below the deepest maize roots did not occur, resulting in significant (P<0.05) suppression of maize root lengths and downward root growth. Maximum root length densities for both species occurred at the top of the soil profile, reaching 1.1–1.7 cm cm^-3 for G. robusta, but only 0.5 cm cm^-3 for maize grown with trees. Root populations in mixed plots were dominated by G. robusta at all times, all depths and all distances from trees and maize and, thus, there was no spatial separation of the rooting zones of the trees and crop. Competition between G. robusta and maize for soil water stored near the surface was unavoidable, although pruning reduced its impact; complementary use of water by the trees and crop would only have been possible if alternative sources of water were available. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root zone solute dynamics under drip irrigation: A review

Infiltration and subsequent distribution of water and solutes under cropped conditions is strongly dependent on the irrigation method, soil type, crop root distribution, and uptake patterns and rates of water and solutes. This review discusses aspects of soil water and solute dynamics as affected by the irrigation and fertigation methods, in the presence of active plant uptake of water and solutes. Fertigation with poor quality water can lead to accumulation of salts in the root zone to toxic levels, potentially causing deterioration of soil hydraulic and physical properties. The high frequency of application under drip irrigation enables maintenance of salts at tolerable levels within the rooting zone. Plant roots play a major role in soil water and solute dynamics by modifying the water and solute uptake patterns in the rooting zone. Modeling of root uptake of water and solutes is commonly based on incorporating spatial root distribution and root length or density. Other models attempt to construct root architecture. Corn uptake rate and pattern of nitrate nitrogen was determined from field studies of nitrate dynamics under drip irrigation using TDR monitoring. The determined nitrate nitrogen uptake rates are within literature values for corn. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root distribution of poplar at varying densities on pastoral hill country

Spaced poplar (Populus spp.) trees are used widely in New Zealand for soil conservation on erodible pastoral hill country. Their root distribution in this environment, and factors that affect it, are poorly understood. Robust recommendations on effective tree spacing depend on knowledge of root systems. This study determined the effect of tree density, position between trees, and soil depth (0–90 cm) on root number, root diameter distribution, root area ratio (RAR), and cross sectional area per root for young trees on slopes. Data were collected for lateral roots using trenches. Greater than 80% of roots were < 5 mm diameter and root attributes were highest in shallow soil. Trees at 770 stems per hectare (sph) had 3–12 times more roots and 3–9 times greater RAR than those at densities of ≤ 237 sph, representative of most tree-pasture systems. Mean cross sectional area per root was similar across densities. Positions close to trees had twice as many roots (46 vs. 23/m²) and RAR (109 vs. 52 mm²/m²) as positions midway between trees. The study provided quantitative understanding of variation in root distribution with tree density and information useful for supporting and strengthening recommendations on densities for effective erosion control.     

Root respiration during grain filling in sunflower: The effects of water stress

Instantaneous rates of (soil + root) respiration were measured periodically during grain filling in sunflower crops that were i) irrigated at weekly intervals and ii) subjected to water stress for the last 25 days of the 40-day grain filling period. Daily (soil + root) respiration was calculated using instantaneous respiration rates, an empirically determined temperature response function, and diurnal records of soil temperature. Daily soil respiration was estimated using empirically determined functions linking soil respiration to soil temperature and water content. Between anthesis and maturity, daily root respiration of the irrigated crop dropped by about one half from ca. 1.8 g C m^-2 d^-1, exhibiting a strong association with daily crop gross photosynthesis. Water stress brought about a rapid decrease in root respiration, which fell to about 0.1 g C m^-2 d^-1 at maturity. Root respiration during grain filling was 46 and 30 g C m^-2 for irrigated and stressed crops, respectively.     

Root growth and water uptake during water deficit and recovering in wheat

Root growth and soil water content were measured in a field experiment with wheat subjected to two periods of water deficit. The first period was induced early in the season between the early vegetative stage (22 DAS) and late terminal spikelet (50 DAS), the second period at mid-season between terminal spikelet (42 DAS) and anthesis (74 DAS). Total root growth was reduced under water deficit by a reduction in the top 30 cm, while the root system continued to grow in the deeper soil profile between 30 and 60 cm. Shortly after rewatering, the growth pattern reverted to fastest root growth rates in the shallow soil layers. In relative terms, the total root system increased in relation to the above ground dry matter under water shortage. The early-, the mid-season water deficit treatments, and the control treatment had total root length of 27.4, 19.4 and 30.6 km m^-2, respectively, about 2 wk before maturity. Evapotranspiration declined under water deficit, but water uptake in deeper layers increased. Water uptake per unit root length was reduced with water deficit and was still low shortly after rewatering. Remarkable was the increase in water uptake at 2–3 weeks after rewatering, both deficit treatments exceeded the control by almost 100%. This increase in water uptake followed the burst of new root growth in the upper regions of the soil. However, water uptake rates subsequently declined towards maturity, being between 0.15 L km^-1 d^-1 and 0.17 L km^-1 d^-1 for the early and mid-season water deficit treatments, slightly higher than the control, 0.12 L km^-1 d^-1. The results showed that the crop subjected to early water deficit could compensate for some of the reductions in root growth during subsequent rewatering, but the impact of the mid-season water deficit treatment was more severe and permanent.     

Root foraging strategies and soil patchiness in a humid savanna

In Lamto (Côte d'Ivoire), the savanna is a patchy environment as far as soil is concerned: tree clumps and termite mounds lead to higher nutrient contents than in the surrounding savanna. Mature Borassus aethiopum (Mart.) specimens are tall palm trees dominating the community, with aerial parts located out of these nutrient-rich patches.Palm root densities were compared under tree clumps and in the surrounding savanna, and were also sampled along transects between palm trees and nutrient-rich patches (two clumps and one mound). Palm root densities were far higher (up to 10 times) in the nitrogen-rich soil of both clumps and termite mounds than in the surrounding savanna. Evidence is given that palm trees are able to extend their root system as far as 20 m towards these nutrient-rich patches where they proliferate. These results point out a particular root foraging strategy, which is one of the first known for a woody perennial. They also provide new insights for understanding nitrogen cycling and savannas high rate of primary production.     

Relationship between Root Growth of Potato,Root Diffusate Production,and Hatching of Globodera rostochiensis

Hatching response of Globodera rostochiensis in potato root diffusate (PRD) collected by soaking individual potato, Solanum tuberosum, root systems in water for 2 hours was used to assess the relationship between root growth and PRD production. Resistant potato cultivars Hudson and Rosa were used as test plants. Maximum hatch occurred in PRD collected 3 weeks after plant emergence (AE) in the greenhouse, and declined after this time. Hatch was positively correlated with increased root weight only during the first 3 weeks AE. Hudson PRD was consistently more active than Rosa PRD in stimulating hatch, except when adjusted for root weight. Although the results indicated that cells at the root tip produced a more active PRD than cells located elsewhere, PRD appeared to be produced along the entire root. Differences in time length of the vegetative growth phase, extent of root growth, and volume of roots, rather than the production of a more active PRD per se, may explain why Hudson is more effective than Rosa in reducing G. rostochiensis population densities in soil.     

Root attributes affecting water uptake of rice (Oryza sativa) under drought

Lowland rice roots have a unique physiological response to drought because of their adaptation to flooded soil. Rice root attributes that facilitate growth under flooded conditions may affect rice response to drought, but the relative roles of root structural and functional characteristics for water uptake under drought in rice are not known. Morphological, anatomical, biochemical, and molecular attributes of soil-grown rice roots were measured to investigate the genotypic variability and genotype×environment interactions of water uptake under variable soil water regimes. Drought-resistant genotypes had the lowest night-time bleeding rates of sap from the root system in the field. Diurnal fluctuation predominated as the strongest source of variation for bleeding rates in the field and root hydraulic conductivity (Lp (r)) in the greenhouse, and was related to expression trends of various PIP and TIP aquaporins. Root anatomy was generally more responsive to drought treatments in drought-resistant genotypes. Suberization and compaction of sclerenchyma layer cells decreased under drought, whereas suberization of the endodermis increased, suggesting differential roles of these two cell layers for the retention of oxygen under flooded conditions (sclerenchyma layer) and retention of water under drought (endodermis). The results of this study point to the genetic variability in responsiveness to drought of rice roots in terms of morphology, anatomy, and function.     

Root hair length and rhizosheath mass depend on soil porosity,strength and water content in barley genotypes

Selecting plants with improved root hair growth is a key strategy for improving phosphorus-uptake efficiency in agriculture. While significant inter- and intra-specific variation is reported for root hair length, it is not known whether these phenotypic differences are exhibited under conditions that are known to affect root hair elongation. This work investigates the effect of soil strength, soil water content (SWC) and soil particle size (SPS) on the root hair length of different root hair genotypes of barley. The root hair and rhizosheath development of five root hair genotypes of barley (Hordeum vulgare L.) was compared in soils with penetrometer resistances ranging from 0.03 to 4.45 MPa (dry bulk densities 1.2–1.7 g cm^?3). A “short” (SRH) and “long” root hair (LRH) genotype was selected to further investigate whether differentiation of these genotypes was related to SWC or SPS when grown in washed graded sand. In low-strength soil (<1.43 MPa), root hairs of the LRH genotype were on average 25 % longer than that of the SRH genotype. In high-strength soil, root hair length of the LRH genotype was shorter than that in low-strength soil and did not differ from that of the SRH genotype. Root hairs were shorter in wetter soils or soils with smaller particles, and again SRH and LRH did not differ in hair length. Longer root hairs were generally, but not always, associated with larger rhizosheaths, suggesting that mucilage adhesion was also important. The root hair growth of barley was found to be highly responsive to soil properties and this impacted on the expression of phenotypic differences in root hair length. While root hairs are an important trait for phosphorus acquisition in dense soils, the results highlight the importance of selecting multiple and potentially robust root traits to improve resource acquisition in agricultural systems.     

Root development and configuration in intensively managed radiata pine plantations

Summary In south-east Australia, where radiata pine (Pinus radiata D. Don) is grown on sandy soils low in nutrients and short of water, early establishment, and rapid growth to canopy closure lead to increased productivity. At this stage demands for nutrients and water are high, and trees respond vigorously to silvicultural inputs.For several months after transplanting in winter roots are confined within a narrow planting wedge, low temperature restricts new root growth and slows recovery from water stress in plants. From spring, depending upon the configuration and vigour of the roots transplanted, lateral roots extend radially throughout the soil.Although there were small decreases in concentration of roots radially from the stems of very young trees, such spatial differences disappeared between ages 2 and 3, so that rooting density was independent of distance from the stem. The pattern of vertical distribution of lateral roots was not influenced by age and 80–90% of the lateral roots were within the top 30 cm soil. Roots developed rapidly as the trees grew towards canopy closure, but in general the rooting densities of these pines are among the lowest reported for plants. In rapidly growing trees approaching canopy closure, the secondary thickening of the lateral roots was sufficient to double the weight of roots without altering root length.Knowledge about root growth and root configuration during the early phase of plantation development will assist management decisions where intensive silviculture is practiced, and hence ensure the most efficient use of nutrients and water.     

Root characteristics of representative Mediterranean plant species and their erosion-reducing potential during concentrated runoff

Gully erosion is an important soil degradation process in Mediterranean environments. Revegetation strategies for erosion control rely in most cases on the effects of the above-ground biomass on reducing water erosion rates, whereas the role of the below-ground biomass is often neglected. In a Mediterranean context, the above-ground biomass can temporally disappear because of fire or overgrazing and when concentrated flow erosion occurs, roots can play an important role in controlling soil erosion rates. Unfortunately, information on root characteristics of Mediterranean plants, growing on semi-natural lands, and their effects on the topsoil resistance to concentrated flow erosion is lacking. Therefore, typical Mediterranean grass, herb, reed, shrub and tree root systems of plants growing in habitats that are prone to concentrated flow erosion (i.e. in ephemeral channels, abandoned fields and steep badland slopes) are examined and their erosion-reducing potential was evaluated. Root density (RD), root length density (RLD) and root diameters are measured for 26 typical Mediterranean plant species. RD values and root diameter distribution within the upper 0.10–0.90 m of the soil profile are then transformed into relative soil detachment rates using an empirical relationship in order to predict the erosion-reducing effect of root systems during concentrated runoff. Comparing the erosion-reducing potential of different plant species allows ranking them according to their effectiveness in preventing or reducing soil erosion rates by concentrated flow. RD in the 0.10 m thick topsoil ranges between 0.13 kg m⁻³ for Bromus rubens (L.) and 19.77 kg m⁻³ for Lygeum spartum (L.), whereas RLD ranges between 0.01 km m⁻³ for Nerium oleander (L.) and 120.43 km m⁻³ for Avenula bromoides ((Gouan) H. Scholz.) Relative soil detachment rates, compared to bare soils, range between 0.3 × 10^-12 and 0.7 for the 0.10 m thick topsoil. The results show that grasses such as Helictotrichon filifolium ((Lag.) Henrard), Piptatherum miliaceum ((L.) Coss.), Juncus acutus (L.), Avenula bromoides ((Gouan) H. Scholz), Lygeum spartum (L.) and Brachypodium retusum ((Pers.) Beauv.) have the highest potential to reduce soil erosion rates by concentrated flow in the 0–0.1 m topsoil. But also shrubs such as Anthyllis cytisoides (L.) and Tamarix canariensis (Willd.), having high root densities in the topsoil, can reduce erosion rates drastically. Among the species growing in channels, Juncus acutus (L.) has the highest erosion reducing potential, whereas Phragmites australis (Cav.) is the least effective. On abandoned fields, Avenula bromoides ((Gouan) H. Scholz) and Plantago albicans (L.) are the most effective species in reducing concentrated flow erosion rates, while Thymelaea hirsuta (L. (Endl.)) and Bromus rubens (L.) perform the worst. On steep badland slopes, Helictotrichon filifolium ((Lag.) Henrard) and Anthyllis cytisoides (L.) perform the best in the analysis of erosion reducing potential, while Ononis tridentata (L.) is the least effective species. These findings have implications for ecological restoration and management of erosion-prone slopes.     

Root growth of triticale and barley grown for grain or for forage-plus-grain in a Mediterranean climate

Root systems of one triticale (× triticosecale Witt.) and one barley (Hordeum vulgare L.) cultivar grown for grain or for the dual purpose of winter forage-plus-grain were studied in a Mediterranean climate (Granada, Spain). The aim was to assess the effect of winter forage removal on root systems and to improve the knowledge of cereal root systems under Mediterranean conditions in relation to soil water use. After the forage was removed by clipping at the end of the winter period, cereal roots were shallower and lower in length densities, compared to the unclipped treatment. The largest differences occurred during the clipping-anthesis period and in the upper soil layers. At the end of the life cycle, the differences between the two systems regarding depth, density and dry matter of roots were small or nil. Moreover, there were no differences in total water use between clipped and unclipped cereals.Under both production systems (grain and forage plus grain), cereals demonstrated variable downward root extension (0.9 to 1.8 m) as a response to the wetting depth. Triticale roots continued growing after anthesis, especially in the deeper soil layers. In spite of that, root systems were not able to extract a notable amount of residual water (25 to 50 mm) apparently available from the subsoil. In semi-arid Mediterranean drylands, cereal root systems with greater phenotypic plasticity (deeper or larger in the subsoil) in response to subsoil water should be of interest in wetter areas or seasons. This does not necessarily imply a larger root system, but rather a root growth pattern with greater root growth in the subsoil.     

Root respiration in citrus acclimates to temperature and slows during drought

Citrus seedlings were grown in soil columns in which the root system was hydraulically separated into two equal layers; this enabled us to maintain roots in the upper layer without water for 110 d. The columns were placed into waterbaths modified so that soil temperatures in the top layer could be maintained at 25°C or at 35°C, while temperature in the bottom layer was maintained at 25°C. We hypothesized that, if citrus plants were grown in dry soil for an extended period, root mortality would increase if the cost of maintaining the roots was increased by elevating the soil temperature. However, during the drought period we did not observe any root mortality, even at the higher soil temperature. Moreover, we did not find that root respiration was increased by prolonged exposure to drought and higher soil temperature. We did find that root respiration rates slowed in dry soil. Furthermore, when the soil columns were switched from one temperature treatment to another, root respiration rates in wet soil rapidly increased when moved to a higher temperature or rapidly decreased when moved to a lower temperature. But after only 4 d, respiration rates returned to their original level; root respiration in dry soil was not affected by either short-or long-term shifts in soil temperature. Root respiration in citrus appears to acclimate rapidly to changes in soil temperature.     

Root effects on soil water and hydraulic properties

Plants can affect soil moisture and the soil hydraulic properties both directly by root water uptake and indirectly by modifying the soil structure. Furthermore, water in plant roots is mostly neglected when studying soil hydraulic properties. In this contribution, we analyze effects of the moisture content inside roots as compared to bulk soil moisture contents and speculate on implications of non-capillary-bound root water for determination of soil moisture and calibration of soil hydraulic properties. In a field crop of maize (Zea mays) of 75 cm row spacing, we sampled the total soil volumes of 0.7 m × 0.4 m and 0.3 m deep plots at the time of tasseling. For each of the 84 soil cubes of 10 cm edge length, root mass and length as well as moisture content and soil bulk density were determined. Roots were separated in 3 size classes for which a mean root porosity of 0.82 was obtained from the relation between root dry mass density and root bulk density using pycnometers. The spatially distributed fractions of root water contents were compared with those of the water in capillary pores of the soil matrix. Water inside roots was mostly below 2–5% of total soil water content; however, locally near the plant rows it was up to 20%. The results suggest that soil moisture in roots should be separately considered. Upon drying, the relation between the soil and root water may change towards water remaining in roots. Relations depend especially on soil water retention properties, growth stages, and root distributions. Gravimetric soil water content measurement could be misleading and TDR probes providing an integrated signal are difficult to interpret. Root effects should be more intensively studied for improved field soil water balance calculations. Presented at the International Conference on Bioclimatology and Natural Hazards, Pol’ana nad Detvou, Slovakia, 17–20 September 2007.     

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。     

Root Respiration for Agave deserti: Influence of Temperature, Water Status and Root Age on Daily Patterns

Root respiration, measured as CO₂ efflux, was studied for asucculent perennial from the Sonoran Desert, Agave deserti,with a new technique using individual, attached roots. The dailypatterns of root respiration closely followed the daily patternsof root temperature for both established roots and rain roots,with higher rates during the day when root temperature averaged27?C and lower rates at night when root temperature averaged17?C. When root temperature was raised from 5?C to 40?C, rootrespiration increased about 7-fold; from 45 ?C to 55 ?C, rootrespiration decreased about 2-fold, except for old establishedroots. Root respiration per unit dry weight for both root typesdecreased with age, the initial decrease being greater for rainroots than for established roots. Root respiration rates forrain roots were reduced to zero at a soil water potential (_soil)of –0.9 MPa and did not recover upon rewatering. Upondrying, root respiration rates for established roots were maintainedat about 12% of maximum, even when _soil fell to –1.6 MPa,and fully recovered 1.5 d after rewatering the soil. Such responsesof rain and established roots must be taken into account whenassessing the carbon costs for the root system. Key words: Agave deserti, CO₂ exchange, root respiration, temperature, soil water potential