Growth of a root system described as diffusion. II. Numerical model and application

In simulation models for water movement and nutrient transport, uptake of water and nutrients by roots forms an essential part. As roots are spatially distributed, prediction of root growth and root distribution is crucial for modelling water and nutrient uptake. In a preceding paper, De Willigen et al. (2002; Plant and Soil 240, 225–234) presented an analytical solution for describing root length density distribution as a diffusion-type process. In the current paper, we present a numerical model that does the same, but which is more flexible with respect to where root input can occur. We show that the diffusion-type root growth model can describe well observed rooting patterns. We used rooting patterns for different types of crops: maize, gladiolus, eastern white cedar, and tomato. For maize, we used data for two different types of fertiliser application: broadcast and row application. In case of row application, roots extend more vertically than horizontally with respect to the broadcast application situation. This is reflected in a larger ratio of diffusion coefficients in vertical versus horizontal direction. For tomato, we considered tomatoes grown on an artificial rooting medium, i.e. rockwool. We have shown that, in principle, the model can be extended by including reduction functions on the diffusion coefficient in order to account for environmental conditions.     

Seasonal dynamics of fine root biomass,root length density,specific root length,and soil resource availability in a Larix gmelinii plantation

Fine root tumover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors.Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past,our understanding of it remains limited.This is because the dynamics processes associated with soil resources availability are still poorly understood.Soil moisture,temperature,and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level.In temperate forest ecosystems,seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground.Therefore,fine root biomass,root length density(RLD)and specific root length(SRL)vary during the growing season.Studying seasonal changes of fine root biomass,RLD,and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover.The objective of this study was to understand whether seasonal variations of fine root biomass,RLD and SRL were associated with soil resource availability,such as moisture,temperature,and nitrogen,and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation.We used a soil coring method to obtain fine root samples(≤2 mm in diameter)every month from Mav to October in 2002 from a 17-year-old L.gmelinii plantation in Maoershan Experiment Station,Northeast Forestry University,China.Seventy-two soil cores(inside diameter 60 mm;depth intervals:0-10 cm,10-20 cm,20-30 cm)were sampled randomly from three replicates 25 m×30 m plots to estimate fine root biomass(live and dead),and calculate RLD and SRL.Soil moisture,temperature,and nitrogen(ammonia and nitrates)at three depth intervals were also analyzed in these plots.Results showed that the average standing fine root biomass(live (32.2 g.m-2.a-1)in the middle(10-20 cm)and deep layer (20-30cm),respectively.Live and dead fine root biomass was the highest from May to July and in September,but lower in August and October.The live fine root biomass decreased and dead biomass increased during the growing soil layer.RLD and SRL in May were the highestthe other months,and RLD was the lowest in Septemberdynamics of fine root biomass,RLD,and SRL showed a close relationship with changes in soil moisture,temperature,and nitrogen availability.To a lesser extent,the temperature could be determined by regression analysis.Fine roots in the upper soil layer have a function of absorbing moisture and nutrients,while the main function of deeper soil may be moisture uptake rather than nutrient acquisition.Therefore,carbon allocation to roots in the upper soil layer and deeper soil layer was different.Multiple regression analysis showed that variation in soil resource availability could explain 71-73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass.These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability,which resulted in an increased allocation of carbohydrate to these roots,but a lower allocation of carbohydrate to those in soil with lower resource availability.     

Tillage and N fertilization effects on maize root growth and root:shoot ratio

Two methods for estimating the size of the maize (Zea mays l.) root system from soil cores taken in the field were compared. The spatially weighed block method of estimation accounted for variation in root density by using 18 samples per plant which varied in distance from plant and soil depth. This method was compared to an estimation which averaged all of the 18 samples together. Both methods gave surprisingly similar estimates for total root growth. Increased root growth in the surface soil layers, due to tillage and N fertilization, did not impact on the estimation of total root growth. Total root length remained unchanged or increased with N fertilization, while root weight remained the same or decreased. Root mass per length decreased with N fertilization. The estimated size of the root system was used to calculate root:shoot weight ratios. The largest root:shoot ratio was found in the vegetative stage and decreased throughout the rest of the season. In this field experiment, the estimated size of the root system at 8 weeks after planting was not significantly different from the size at silking or harvest. Nitrogen fertilization significantly decreased the root:shoot weight ratio. However, tillage did not significantly change the ratio.     

Effect of cyclic wetting and drying of a soil on root hair growth of maize roots

The objective of this study was to investigate the effect of cyclic soil wetting and drying on maize (Zea mays L.) root hair growth. Three soils, Chalmers silty clay loam (Typic Haplaquolls), Raub silt loam (Aquic Argiudolls) and Aubbeenaubbee sandy loam (Aric Ochraqualfs) and two soil moisture contents, −175 (M₀) and −7.5 kPa (M₁), were used to study root hair growth in a controlled-climate chamber. Increasing soil moisture after 7d from M₀ and M₁ resulted in a cessation of root hair growth behind the root cap while drying the soil after 7d from M₁ and M₀ promoted root hair growth on new but not old or existing roots. By maintaining liquid continuity under cyclic wetting and drying of a soil, root hairs may be of far greater significance to the nutrition of the plant than originally thought. Journal Paper No. 11023, Purdue Univ. Agric. Exp. Stn., W. Lafayette, IN 47907. Contribution from the Dep. of Agron.     

Seasonal dynamics of fine root biomass, root length density, specific root length, and soil resource availability in a Larix gmelinii plantation

Fine root turnover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors. Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past, our understanding of it remains limited. This is because the dynamics processes associated with soil resources availability are still poorly understood. Soil moisture, temperature, and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level. In temperate forest ecosystems, seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground. Therefore, fine root biomass, root length density (RLD) and specific root length (SRL) vary during the growing season. Studying seasonal changes of fine root biomass, RLD, and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover. The objective of this study was to understand whether seasonal variations of fine root biomass, RLD and SRL were associated with soil resource availability, such as moisture, temperature, and nitrogen, and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation. We used a soil coring method to obtain fine root samples (⩽2 mm in diameter) every month from May to October in 2002 from a 17-year-old L. gmelinii plantation in Maoershan Experiment Station, Northeast Forestry University, China. Seventy-two soil cores (inside diameter 60 mm; depth intervals: 0–10 cm, 10–20 cm, 20–30 cm) were sampled randomly from three replicates 25 m × 30 m plots to estimate fine root biomass (live and dead), and calculate RLD and SRL. Soil moisture, temperature, and nitrogen (ammonia and nitrates) at three depth intervals were also analyzed in these plots. Results showed that the average standing fine root biomass (live and dead) was 189.1 g·m⁻²·a⁻¹, 50% (95.4 g·m⁻²·a⁻¹) in the surface soil layer (0–10 cm), 33% (61.5 g·m⁻²·a⁻¹), 17% (32.2 g·m⁻²·a⁻¹) in the middle (10–20 cm) and deep layer (20–30cm), respectively. Live and dead fine root biomass was the highest from May to July and in September, but lower in August and October. The live fine root biomass decreased and dead biomass increased during the growing season. Mean RLD (7,411.56 m·m⁻³·a⁻¹) and SRL (10.83 m·g⁻¹·a⁻¹) in the surface layer were higher than RLD (1 474.68 m·m⁻³·a⁻¹) and SRL (8.56 m·g⁻¹·a⁻¹) in the deep soil layer. RLD and SRL in May were the highest (10 621.45 m·m⁻³ and 14.83m·g⁻¹) compared with those in the other months, and RLD was the lowest in September (2 198.20 m·m⁻³) and SRL in October (3.77 m·g⁻¹). Seasonal dynamics of fine root biomass, RLD, and SRL showed a close relationship with changes in soil moisture, temperature, and nitrogen availability. To a lesser extent, the temperature could be determined by regression analysis. Fine roots in the upper soil layer have a function of absorbing moisture and nutrients, while the main function of deeper soil may be moisture uptake rather than nutrient acquisition. Therefore, carbon allocation to roots in the upper soil layer and deeper soil layer was different. Multiple regression analysis showed that variation in soil resource availability could explain 71–73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass. These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability, which resulted in an increased allocation of carbohydrate to these roots, but a lower allocation of carbohydrate to those in soil with lower resource availability. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(3): 403–410 [译自: 植物生态学报, 2005, 29(3): 403–410]     

Modelling the branching growth fractal pattern of the maize root system

Using the technique of L-systems, a growth model of the maize root system is developed. From the observation of the root systems developed under various soil density in eight root boxes, a spatial hierarchy of growth rules was extracted. The rules were divided into three categories: a meta-rule for describing features of an entire root system, a branching growth rule and a tip elongation rule. Some variations in the entire features of the root system, such as the outline and the root distribution, were confirmed by observation, and then the respective meta-rules were re-defined. The branching properties of first- and second-order lateral roots were statistically almost equal in the observations, and this lead us to set up a single stochastic branching growth rule. Tip elongation movement was not observed here; its rule had to be assumed by reference to data in the literature. A single set of branching growth and tip elongation rules were coupled with the respective meta-rules corresponding to the root samples observed, where a small scale rule was loosely governed by a large scale rule. Computer simulations offered optimized drawings of the observed root systems, and they also reproduced a typical anisotropic power distribution of roots similar to those observed.     

Significance of temperature and precipitation for maize root distribution in the field

Measurements of maize (Zea mays L.) root distribution with depth in the soil for nine years in a 11-year period revealed significantly different distribution patterns. Weather variations were expected to be related to the amount of roots found in each of the five 15-cm soil layers. The objective of this study was to attempt to explain root distribution in the field on the basis of precipitation and temperature data for the nine growing seasons. Growing degree days (GDD), accumulated in daily increments from planting to silking, were used to describe temperature effects. Correlations were calculated for weekly time increments of GDD versus root length densities at silking in all soil layers. Root length density below 30 cm was correlated (P=0.05) with GDD for two weeks following planting, whereas no relation was found between GDD and root length density in the topsoil. Amount of precipitation was accumulated in weekly increments from silking to planting and correlated with root length density in the soil layers at silking. This procedure evaluated the relation between precipitation and root growth during the vegetative growth period. Root length density in the 0 to 15 cm layer was found to be related significantly (P=0.05) to precipitation. The period 3 weeks prior silking gave the highest correlation coefficient (r=0.79). Journal Paper no. 10,629. Purdue Univ. Agric. Exp. Stn., W. Lafayette, IN 47907. Contribution from the Dep. of Agronomy. The research was supported in part by BARD, United States-Israel Binational Agricultural Research and Development Fund, and Deutsche Forschungsgemeinschaft.     

Comparison of tomato root distributions by minirhizotron and destructive sampling

Calibration of minirhizotron data against root length density (RLD) was carried out in a field trial where three drip irrigation depths: surface (R0) and subsurface, 0.20 m (RI) and 0.40 m depth (RII) and two processing tomato cultivars: `Brigade' (CI) and `H3044' (CII) were imposed. For each treatment three minirhizotron tubes were located at 10, 37.5 and 75 cm of the way from one plant row to the next. Roots intersecting the minirizotrons walls were expressed as root length intensity (L _a) and number of roots per unit of minirhizotron wall area (N _ra). Root length density (RLD) was calculated from core samples taken for each minirhizotron tube at two locations: near the top of the minirhizotron (BI) and 15 cm apart from it, facing the minirhizotron wall opposite the plant row (BII). Minirhizotron data were regressed against RLD obtained at BI and BII and with their respective means. The results show that for all the situations studied, better correlations were obtained when RLD was regressed with L _a than with N _ra. Also was evident that the relationship between L _a and RLD was strongly influenced by the location of soil coring. RLD was correlated with L _a trough linear and cubic equations, having the last ones higher determination coefficients. For instance at 10 cm from the plant row when values from the top layer (0–40 cm) were analysed separately, L _a was significantly regressed with RLD measured at BII and described by the equations: RLD = 0.5448 + 0.0071 L _a (R ² = 0.51) and RLD = 0.4823 + 0.0074L _a + 8×10^–5 L _a ² – 5×10^–7 L _a ³ (R ² = 0.61). Under the 40 cm depth the highest coefficients of determination for the linear and cubic equations were respectively 0.47 and 0.88, found when L _a was regressed with RLD measured at BI. For minirhizotrons located at 75 cm from the plant row and for location BI it was possible to analyse jointly data from all depths with coefficients of determination of 0.45 and 0.59 for the linear and cubic equations respectively.     

Nutrient uptake relationship to root characteristics of rice

Data on root parameters and distribution are important for an improved understanding of the factors influencing nutrient uptake by a crop. Therefore, a study was conducted on a Crowley silt loam at the Rice Research and Extension Center near Stuttgart, Arkansas to measure root growth and N, P and K uptake by three rice (Oryza sativa L.) cultivars at active tillering (36 days after emergence (DAE)), maximum tillering (41 DAE), 1.25 cm internode elongation (55 DAE), booting (77 DAE) and heading (88 DAE). Soil-root core samples were taken to a depth of 40 cm after plant samples were removed, sectioned into 5 cm intervals, roots were washed from soil and root lengths, dry weights and radii were measured. Root parameters were significantly affected by the soil depth × growth stage interaction. In addition, only root radius was affected by cultivar. At the 0- to 5-cm soil depth, root length density ranged from 38 to 93 cm cm^-3 throughout the growing season and decreased with depth to about 2 cm cm^-3 in the 35- to 40-cm depth increment. The increase in root length measured with each succeeding growth stage in each soil horizon also resulted in increased root surface area, hence providing more exposed area for nutrient uptake. About 90% of the total root length was found in the 0- to 20-cm soil depth throughout the season. Average root radius measured in the 0- to 5-cm and 35- to 40-cm depth increments ranged from 0.012 to 0.013 cm and 0.004 to 0.005 cm, respectively throughout the season. Total nutrient uptake by rice differed among cultivars only during vegetative growth. Differences in total nutrient uptake among the cultivars in the field appear to be related to absorption kinetics of the cultivars measured in a growth chamber study. Published with permission of the Arkansas Agricultural Experiment Station.     

Root growth and distribution of two short-season rice genotypes

Root growth dynamics of lowland rice (Oryza sativa L.) throughout the growing season are poorly understood. A field experiment was conducted in 1987 to compare root growth and distribution of two rice genotypes at two Arkansas locations on soils with different physical and chemical properties. Two genotypes, Bond and an experimental line (RU8701084), were grown on a Captina silt loam (Typic Fragiudults) at Fayetteville, AR, and on a Crowley silt loam (Typic Albaqualfs) near Stuttgart, Ar. Plots contained minirhizotrons oriented at a 45° angle and extended 55 cm (Captina) and 40 cm (Crowley) vertical to the soil surface. Root measurements were taken several times during the season at specific growth stages. Plant height and tiller number were taken at 9 dates at Fayetteville up to physiological maturity. In general, root length (RL) and root length density (RLD) were greater on the Captina soil. Genotypes at both locations reached maximum root growth rates between active tillering and panicle initiation (PI) and maximum RL by early reproduction. Total RL were similar between genotypes on the Captina. However on the Crowley, the mean RL for Bond between the period of early booting and flood removal was an average of 54% greater than for RU8701084. During early reproductive growth at both locations RL plateaued, but then declined during the grain filling process. There was a trend for RU8701084 to contain a greater percentage of its total RL in the top 20 and 10 cm of soil on the Captina and Crowley, respectively, while Bond tended to be a deeper rooted genotype. Bond had a greater RLD at the 20–30 cm depth increment on the Crowley, which contributed to the greater RL. Less than 15% of the total RL of either genotype was measured below 30 cm on the Crowley. In contrast, nearly 25% of the total RL was found at the 30–40 cm depth increment on the Captina. Results showed that rice root growth varied between soils, that root distribution patterns differed between genotypes, and that patterns of root growth changed with changes in plant development.     

Cultivar and planting date effects on soybean root growth

To avoid late summer drought, soybean [Gylcine max (L) Merrill] producers in many southern and border states of the USA modify their cropping systems. Options include use of unadapted cultivars and changing planting dates. Because root function is important to plant health and yield, this study was conducted to determine if planting date and soybean cultivar affect root growth and distribution. Seeds of one cultivar from each of four maturity groups (MG III, IV, V, and VI) were sown in mid-April, mid-May, and mid-June in 1992 and 1993 on a Tiptonville silt loam near Portageville, MO. Root observations were performed 30 and 60 days after emergence (DAE) using a minirhizotron system. Cultivars differed for root length density (RLD) only in the 15 to 28 cm depth in 1992 and in the 15 to 28 cm and 29 to 42 cm depths in 1993, but differences were not related to maturity classification of cultivar. Average RLD was 1.02 cm^–3 for MG III and IV cultivars and 1.21 cm cm^–3 for MG V and VI cultivars. Average RLD for the mid-June planting date was 1.65 cm cm^–3 but only 0.73 cm cm^–3 for the mid-April planting date. An increase in RLD between 30 and 60 DAE occurred at all soil depths. For both years, MG V and VI cultivars produced higher yields than the MG III cultivars. Earlier than normal planting dates inhibited early root growth, but did not reduce yield. Cultivars differed only slightly for the rooting characteristics measured in this study. These rooting characteristics may not be important criteria for cultivar selection.Abbreviations MG maturity group - VCR videocassette recorder - DAE days after emergence - RLD root length density - CRLD change in root length density Contribution from the Missouri Agric. Exp. Station Journal Series Number 12, 153Contribution from the Missouri Agric. Exp. Station Journal Series Number 12, 153     

A growth chamber for idealized studies of seedling root growth dynamics and structure

A root growth chamber is described which allows seedling root growth dynamics and structure to be monitored continuously under a variety of conditions for several weeks. The chamber consists of two cells with inner dimensions 18×20×0.12 cm. To simulate the soil matrix, each cell was filled with spherical glass beads of 0.1 cm diameter. Given the 0.12 cm width of each cell, the glass bead matrix was approximately one bead layer thick. Roots were therefore grown in a quasi -two-dimensional and transparent environment. This enabled root images of high spatial and temporal resolution to be collected and analysed quantitatively using standard image analysis techniques. The chamber was constructed such that the root environment could be manipulated with regard to nutrient distribution, `soil' matrix structure and other perturbations to the system. Preliminary experiments of the growth dynamics of lentil roots (Lens culinaris L. cv. Verte du Puy) in the chamber were conducted. The majority of the primary and lateral roots followed a similar growth pattern with high growth rates between days 5 and 9 and days 14 and 18 separated by a period of low growth rate between days 10 and 12 after seeding in the chamber. Thus, primary and lateral root growth was to a certain extent synchronized. Lateral roots developed after 3 to 8 days on the outer curve (convex side) of the primary root. The roots shared many of the characteristics of roots developed in three-dimensional systems indicating that the chamber did not induce artificial root behaviour. Thus, the idealized and quantitative studies that can be conducted in the chamber may enable many aspects of the complex interactions between the root system and environment to be studied.     

Differences in root and shoot growth and soil moisture extraction between cotton cultivars in an acid subsoil

Research was conducted to determine if differences in yield and crop growth of field-grown cotton cultivars (Stoneville 825, Deltapine 41, Auburn 56, and Pima S-5) would be related to root length density and end-of-season soil moisture content in an acid soil. Soil core root density differences between cultivars were inconsistent between years. However, normalization of root density on a percentage of total root density basis indicated Stoneville 825 and Pima S-5 had a consistently greater percentage of roots in the acidic subsoil than did Auburn 56 and Deltapine 41. Subsoil moisture remaining at the end of the season was least for Stoneville 825 and greatest for Deltapine 41. Shoot growth parameters of Stoneville 825 were numerically and often significantly greater than Deltapine 41 each year. Auburn 56 was comparable or superior to Stoneville 825 and Pima S-5 in most parameters of shoot growth. Cotton cultivar differences in root length density and implied soil moisture extraction in acidic subsoil may partly explain differences in adaptation by some cultivars to nonirrigated, drought prone, acidic soils.     

Effect of irrigation frequency on root water uptake in sugar beet

A 2-year trial was performed on autumn-sown sugar beet grown in pots in order to study the influence of irrigation frequency on the water used by plants along the soil profile. The outdoor pots, containing one plant each, were 1.3 m high and had circular openings, through which Time Domain Reflectometry (TDR) apparatus wave guides could be inserted. Three irrigation intervals were compared and plants were watered whenever the soil layer explored by roots had lost 30% (SWD₁), 50% (SWD₂) and 70% (SWD₃) of the total available water (TAW). During the irrigation season, the water extracted by the plants from each layer along the soil profile (RWU) was determined by monitoring volumetric soil moisture content (), by TDR. At harvest time, root length density (RLD) along the soil profile was assessed using the Tennant method. The applied irrigation frequencies significantly affected the RWU. With the SWD₃ protocol, irrigation was at longer irrigation intervals (9 days) and watering volumes were as high as 84 mm. In this treatment, the plants lost almost 60% of total water from the lower soil layer (0.6–1.0 m). In treatment SWD₁, the irrigation interval was very short (3 days), and water extraction from 0.0–0.6 m soil depth was 92.0%. In the intermediate treatment, the irrigation interval was 5.5 days and a more uniform water depletion was observed along the root zone, approximately equal between the 0–0.6 and 0.6–1.0 m soil layer. Water extraction of sugar beet plants at the deeper soil layers in response to long irrigation intervals was related to an increase in water uptake efficiency of the deeper younger roots and not to an increase in root length density, which, on the contrary, decreased. This morpho-physiological acclimatization to progressive soil water deficit was coupled with an increase of the root/shoot ratio.     

Specific root length as an indicator of environmental change

Abstract

Specific root length (SRL, m g^?1) is probably the most frequently measured morphological parameter of fine roots. It is believed to characterize economic aspects of the root system and to be indicative of environmental changes. The main objectives of this paper were to review and summarize the published SRL data for different tree species throughout Europe and to assess SRL under varying environmental conditions. Meta-analysis was used to summarize the response of SRL to the following manipulated environmental conditions: fertilization, irrigation, elevated temperature, elevated CO₂, Al-stress, reduced light, heavy metal stress and physical disturbance of soil. SRL was found to be strongly dependent on the fine root classes, i.e. on the ectomycorrhizal short roots (ECM), and on the roots <0.5 mm, <1 mm, <2 mm and 1 – 2 mm in diameter SRL was largest for ECM and decreased with increasing diameter. Changes in soil factors influenced most strongly the SRL of ECM and roots <0.5 mm. The variation in the SRL components, root diameter and root tissue density, and their impact on the SRL value were computed. Meta-analyses showed that SRL decreased significantly under fertilization and Al-stress; it responded negatively to reduced light, elevated temperature and CO_2. We suggest that SRL can be used successfully as an indicator of nutrient availability to trees in experimental conditions.     

Plant-derived smoke solutions stimulate the growth of Lycopersicon esculentum roots in vitro

Aqueous extracts of smoke, derived from Themeda triandra, a fire-climax grass, and Passerina vulgaris, a fynbos plant, stimulated the growth of primary root sections of tomato roots in suspension culture. The optimal dilution for both extracts was 1:2000. Several of the fractions obtained from TLC separation of the Themeda and the Passerina extracts significantly promoted primary root growth. The auxins naphthaleneacetic acid (NAA), indolebutyric acid (IBA) and indoleacetic acid (IAA) were found to stimulate the growth of the primary root axis, with IAA and NAA significantly promoting lateral root number. Similarly, the naturally occurring cytokinins, zeatin and its derivatives (zeatin-O-glucoside; dihydrozeatin and zeatin riboside) stimulated primary root length. Zeatin and dihydrozeatin promoted secondary root growth, but only at very low concentrations.