Simulation model of soil compaction and root growth

Soil compaction is a widespread cause of reduced plant productivity. If the effects of soil compaction on plant growth are to be reproduced in simulation models, then the processes through which compaction reduces root elongation must be expressed mathematically and then tested against experimental data. The mathematical theory by which these processes may be represented is given in the accompanying article. In this article, the behavior of a simulation model based on this theory is tested against data for root growth and soil gas concentration recorded from soil columns of which the middle layers were compacted to different bulk densities. The model was able to reproduce the failure of the root system to penetrate the compacted middle layer within the period of the experiment when bulk density exceeded 1.55 Mg m^-3. The model also reproduced decreases in O₂ concentrations, and increases in CO₂ concentrations, in the atmospheres of the compacted layer and of the uncompacted layer below it as bulk density of the compacted layer increased. The simulated time course of O₂ and nutrient uptake and of O₂ concentrations in the compacted layer at different depths is presented and its consistency with experimental findings is examined. As part of a larger ecosystem model, this model will be useful in estimating site-specific effects of soil compaction on carbon cycling in agroecosystems.     

The effect of soil compaction on growth and P uptake by Trifolium subterraneum: interactions with mycorrhizal colonisation

The effects of vesicular-arbuscular mycorrhizal (VAM) colonisation on phosphorus (P) uptake and growth of clover (Trifolium subterraneum L.) in response to soil compaction were studied in three pot experiments. P uptake and growth of the plants decreased as the bulk density of the soil increased from 1.0 to 1.6 Mg m^-3. The strongest effects of soil compaction on P uptake and plant growth were observed at the highest P application (60 mg kg^-1 soil). The main observation of this study was that at low P application (15 mg kg^-1 soil), P uptake and shoot dry weight of the plants colonised by Glomus intraradices were greater than those of non-mycorrhizal plants at similar levels of compaction of the soil. However, the mycorrhizal growth response decreased proportionately as soil compaction was increased. Decreased total P uptake and shoot dry weight of mycorrhizal clover in compacted soil were attributed to the reduction in the root length. Soil compaction had no significant effect on the percentage of root length colonised. However, total root length colonised was lower (6.6 m pot^-1) in highly compacted soil than in slightly compacted soil (27.8 m pot^-1). The oxygen content of the soil atmosphere measured shortly before the plants were harvested varied from 0.18 m³m^-3 in slightly compacted soil (1.0 Mg m^-3) to 0.10 m³m^-3 in highly compacted soil (1.6 Mg m^-3).     

Influence of soil compaction on tunnel network construction by the eastern subterranean termite (Isoptera: Rhinotermitidae)

Eastern subterranean termites, Reticulitermes flavipes (Kollar), workers were introduced into arenas containing low, moderate, and high compaction builder's sand (1.05 g/cm3, 1.18 g/cm3 or 1.35 g/cm3 bulk densities, respectively), and they immediately began tunneling. Termites built the tunnel network significantly fastest in soil of low compaction compared with moderately or highly compacted soil. In soil of low compaction, 221.67 +/- 4.73 cm of total tunnel distance was constructed in 1 d compared with only 96 cm of tunneling in highly compacted soil. At 14 d, total tunnel distance averaged 216.83 +/- 4.56 cm in soil of low compaction compared with 169.70 +/- 4.10 and 181.18 +/- 6.13 cm in moderately and highly compacted soil, respectively. Decreases in total tunnel distance between 1 and 14 d were caused by backfilling of seldom-used tunnels. Termites did the majority of tunneling during the first day of introduction into arenas. In soil of low and moderate compaction, termites essentially constructed the entire tunnel network within the first day, only modifying it by backfilling or maintaining tunnels. In highly compacted soil, 53% of the final tunnel network was constructed during the first day, 87% was constructed by the third day, and 97% was constructed by the seventh day. Soil compaction did not affect the number of primary tunnels or the number and diameter of secondary tunnels. The angle between the secondary tunnel and primary tunnel also was not significantly affected by soil compaction. However, the number of secondary tunnels in soil of low compaction (5.89 +/- 0.51) was significantly greater than in moderately (2.74 +/- 0.36) and highly (3.58 +/- 0.59) compacted soils.     

Virus diseases of subterranean clover

Subterranean clover (Trifolium subterraneum) is grown as a pasture legume in several temperate regions of the world where the soils are acidic and infertile, and the rainfall is winter dominant and less than 600 mm annually. It is particularly important in southern Australia where more than 16 million ha have been sown with this species as the pasture legume component. Nine viruses have been recorded infecting subterranean clover in the field. These are alfalfa mosaic, bean yellow mosaic (pea mosaic), beet western yellows, clover yellow vein, cucumber mosaic, pea enation mosaic, soybean dwarf (subterranean clover red leaf), subterranean clover mottle and white clover mosaic. In addition there is an important problem referred to as subterranean clover stunt that was assumed to be caused by a virus but whose aetiology is still unknown. The importance of these diseases is reviewed and details on their epidemiologies are outlined together with details on progress towards their control and some comments on matters worthy of attention in the future. Reference is also made to several exotic viruses known to infect subterranean clover experimentally that could possible cause problems if introduced into Australia.     

Nutrient uptake and growth of barley as affected by soil compaction

A field experiment with different levels of compaction was carried out on a mouldboard ploughed silty clay, with the objective of studying the effects on plant nutrient uptake and growth. Soil from the field was also used in laboratory studies of carbon and nitrogen mineralization, and plant uptake of water and nutrients. In the field, low as well as high bulk densities reduced biomass production and nutrient uptake of barley (Hordeum vulgare L.) compared to intermediate bulk densities, where grain yield was approximately 20% higher. In the beginning of the growing season, the concentration of phosphorus and potassium was lowest in plants grown in the loosest and in the most compacted soil, and suboptimal for plant growth. The uptake of nutrients transported by diffusion was more affected by compaction than for nutrients transported by mass flow. The reasons for lowered uptake in loose compared to moderately compacted soil could be reduced root-to-soil contact, a low diffusion coefficient for nutrients and/or reduced mass transport of water to seed and roots. Differences in plant nutrient concentrations between treatments gradually declined until harvest. Immediately after compaction there was probably oxygen deficiency in the compacted soil since the air-filled porosity was critically low, but as the soil dried out, mechanical resistance to root growth may have become a more important growth-limiting factor. In the laboratory study, severe compaction reduced carbon mineralization and uptake of water and nutrients by roots, and caused denitrification. There were only small differences between loose and moderately compacted soil in carbon mineralization, nitrogen concentration in the soil, uptake of water and nutrients and dry matter yield. The large yield increase due to recompaction in the field was not reproduced in the laboratory. Possible reasons are differences in soil temperature between the field and laboratory, in the sowing and fertilizing methods, the pretreatment of the soil and in the spatial variability of bulk density. It is possible that recompaction is needed only in the uppermost part of the soil, which is the loosest, dries out first, and is where the seed as well as the fertilizer are placed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Soil-Behaviour of Phytophthora clandestina

Investigations were undertaken to study the nature and behaviour of P. clandestina in soil. The pathogen was recovered only from soil sievings 250–499 μm and 500 μm–0.99 mm, containing small root fragments. In soil the introduced inoculum of the fungus was incapable of saprophytically and competitively colonizing the dead cotyledons of subterranean clover used as bait material. Exposure of the inoculum to increasing numbers of microbes by adding greater proportions of nonsterile fields, oil to the growth medium of the plant had no significant effect on survival rate and fresh shoot weight of subterranean clover. However, microbes present in the field soil reduced the severity of root rot of subterranean clover. P. clandestina was, able to spread between 15–30 mm through pasteurized soil within a period of 20 days.     

Increased damage by western flower thrips Frankliniella occidentalis in chrysanthemum intercropped with subterranean clover

Suppressive effects of intercropping on Frankliniella occidentalis (Pergande) infestations have been reported in several crops. However, this study demonstrates that in year-round chrysanthemum, Dendranthema grandiflora Tzvelev, undersowing with subterranean clover, Trifolium subterraneum L., results in an increased thrips feeding damage. In a pot experiment, performed with chrysanthemum plants (cultivars Reagan and Tiger) grown with or without subterranean clover, significantly more leaves with silver and growth damage were found in the chrysanthemum plants with subterranean clover in comparison with the monocropped chrysanthemum plants. Similarly, the degree of deformation of leaf perimeter and leaf surface was higher in the top leaves of the intercropped chrysanthemum plant. In the soil experiment (only performed with Tiger and plants were grown in the soil in the greenhouse) intercropped chrysanthemum plants suffered a higher feeding damage as well. Analysis of the relation between silver or growth damage and the thrips pressure demonstrates that at similar thrips pressure in the intercropped chrysanthemum plants suffered significantly more damage. The higher thrips pressure in the intercropped chrysanthemum only explains the differences in damage partly. Changes in the reaction of chrysanthemum plants to thrips feeding or in the behaviour of the thrips, mediated by the presence of the non-host crop, are discussed. Our explanation is that chrysanthemum plants grown with clover are more susceptible to thrips feeding than monocropped plants. We conclude that undersowing with clover does not contribute to reduce damage by F. occidentalis in year-round chrysanthemum. Also, the influence of crop diversification on a pest cannot be foreseen until the specific characteristics of each individual crop – pest system are studied.     

Subterranean clover decline in permanent pastures in north-eastern Victoria

Experimental sites were established at two locations in north-eastern Victoria to define factors limiting the establishment and growth of Trifolium subterranean L. (subterranean clover). Liming the soil, seed inoculation and fungicide application were used in renovating subterranean clover pasture on two acidic soils (Longwood: brown/grey sandy loam DY 3.14 and Seymour: grey brown light clay DY 3.22, Northcote classification) with mean annual rainfall of 650 mm and 600 mm respectively. Soil acidity, low available soil phosphorus and plant disease were identified as factors limiting clover yield on these soils. Significant yield responses to lime (35–140%) were obtained with subterranean clover at both sites, with corresponding decreases in Al in the 0–10 cm soil horizon. Liming the soil, when combined with seed inoculation, increased the number and effectiveness of root nodules at both sites. Soil P available for plant growth was low at both sites (6.1 and 8.4 μg g⁻¹) resulting in sub-optimal P concentrations in the clover herbage (45 mmol kg⁻¹ at Longwood). Levels of root disease were low but Aphanomyces euteiches and Phytophthora clandestina (causal agents of lateral and tap root rot) were detected frequently on roots. Application of fungicide resulted in higher dry matter yields (p=0.05) at both sites. An assessment of the relative contributions of these limiting factors and the benefits to be obtained from better management would provide a clearer picture of the profitability and sustainability of this farming system.     

Plant nutrition and growth: Basic principles

Soil compaction may restrict shoot growth of sugar beet plants. Roots, however, are the plant organs directly exposed to soil compaction and should therefore be primarily affected. The aim of this study was to determine the influence of mechanical resistance and aeration of compacted soil on root and shoot growth and on phosphorus supply of sugar beet. For this purpose, a silt loam soil was adjusted to bulk densities of 1.30, 1.50 and 1.65 g cm^–3 and water tensions of 300 and 60 hPa. Sugar beet was grown in a growth chamber under constant climatic conditions for 4 weeks. Both, decrease of water tension and increase of bulk density impeded root and shoot growth. In contrast, the P supply of the plants was differently affected. At the same air-filled pore volume, the P concentration of the shoots was reduced by a decrease of soil water tension, but not by an increase of bulk density. Both factors also reduced root length and root hair formation, however, in compacted soil the plants partly substituted for the reduction of root size by increasing the P uptake efficiency per unit of root. Shoot growth decreased when root growth was restricted. Both characteristics were closely related irrespective of the cause of root growth limitation by either compaction or water saturation. It is therefore concluded that shoot growth in both the compacted and the wet soil was regulated by root growth. The main factor impeding root growth in compacted soil was penetration resistance, not soil aeration.FAX no corresponding author: +49551 5056299     

Capacity of biological soil crusts colonized by the lichen <Emphasis Type="Italic">Diploschistes</Emphasis> to metabolize simple phenols

Background and aims

Soil compaction strongly affects water uptake by roots. The aim of the work was to examine soil—plant interactions with focus on the impact of distribution of compacted soil layers on growth and water uptake by wheat roots.

Methods

The growth-chamber experiment was conducted on wheat growth in soil with compacted soil layers. The system for maintaining constant soil water potential and measurement of daily water uptake from variously compacted soil layers was used.

Results

Layered soil compaction differentiated vertical root distribution to higher extent for root length than root mass. The propagation rate of a water extraction front was the highest through layers of moderately compacted soil. The root water uptake rate was on average 67 % higher from moderately than heavily compacted soil layers. Correlations between water uptake and the length of thick roots were increasing with increasing level of soil compaction.

Conclusions

The study shows that root amount, water uptake, propagation of water extraction and shoot growth strongly depend on the existence of compacted layers within soil profile. The negative effects of heavily compacted subsoil layer on water uptake were partly compensated by increased uptake from looser top soil layers and significant contribution of thicker roots in water uptake.     

Effects of phosphorus application and mycorrhizal inoculation on root characteristics of subterranean clover and ryegrass in relation to phosphorus uptake

The effects of phosphorus (P) application and mycorrhizal inoculation on the root characteristics of subterranean clover and ryegrass were examined. Phosphorus application increased total root length, root surface area and root volume of both plant species. In contrast, mycorrhizal infection only affected the root characteristics of subterranean clover. Ryegrass took up more P than non-mycorrhizal subterranean clover at all levels of application. However, mycorrhizal infection only increased P uptake by subterranean clover and there was no difference in P uptake between ryegrass and mycorrhizal subterranean clover at low levels of P application. When the P uptake was expressed on the basis of any of the root characteristics, subterranean clover were superior to ryegrass suggesting that the greater uptake of P by ryegrass is not due to a higher efficiency in absorption of P from soil solution, but rather to a large root system.     

Effects of Gypsophila saponins on bacterial growth kinetics and on selection of subterranean clover rhizosphere bacteria

Plant secondary metabolites, such as saponins, have a considerable impact in agriculture because of their allelopathic effects. They also affect the growth of soil microorganisms, especially fungi. We investigated the influence of saponins on rhizosphere bacteria in vitro and in soil conditions. The effects of gypsophila saponins on the growth kinetics of rhizosphere bacteria were studied by monitoring the absorbance of the cultures in microtiter plates. Gypsophila saponins (1%) increased the lag phase of bacterial growth. The impact of gypsophila saponins on subterranean clover rhizosphere was also investigated in a pot experiment. The addition of gypsophila saponins did not modify clover biomass but significantly increased (twofold with 1% saponins) the weight of adhering soil. The number of culturable heterotrophic bacteria of the clover rhizosphere was not affected by the addition of gypsophila saponins. Nevertheless, the phenotypical characterization of the dominant Gram-negative strains of the clover rhizosphere, using the Biolog system, showed qualitative and quantitative differences induced by 1% saponins. With the addition of saponins, the populations of Chryseomonas spp. and Acinetobacter spp., the two dominant culturable genera of control clover, were no longer detectable or were significantly decreased, while that of Aquaspirillum dispar increased and Aquaspirillum spp. became the major genus. Aquaspirillum dispar and Aquaspirillum spp. were also the dominant rhizosphere bacteria of Gypsophila paniculata, which greatly accumulates these saponins in its roots. These results suggest that saponins may control rhizosphere bacteria in soil through rhizodeposition mechanisms.     

The effect of soil compaction on the saprophytic growth of Rhizoctonia solani

The increase in bare patch of cereals associated with minimum tillage practices prompted an investigation of the relationship between soil compaction and saprophytic growth of Rhizoctonia solani. In soils wetter than 10 kPa there was a greater density of hyphae in compacted than in non-compacted soil. In relatively dry soil, however, there was wider exploration by hyphae in non-compacted than in compacted soil. The implications of these findings for disease management are discussed.     

The fate of nitrogen (15N) released from different plant materials during decomposition under field conditions

Release of N, retention in soil, availability to a subsequent crop and total recovery of N derived from different¹⁵N-labelled plant materials decomposing in soil was investigated in two field experiments. In the first experiment five different plant species (white clover, red clover, subterranean clover, field bean and timothy) and in the second subterranean clover of different maturity (2,3 and 4 months old) were buried in mesh bags in the soil and allowed to decompose for 10 and 4 months, respectively. Most of the N released from the decaying plant materials was retained in the soil (27–46% of input). The subsequent crop (barley) took up 6–25% of input. The uptake correlated with the amount of N released from the decomposing material (r=0.936^*, I experiment). Similar amounts of subterranean clover N were taken up by barley regardless to whether the material was buried in soil in the previous autumn or just before sowing of the crop. At the end of the experiments, the total recovery of the introduced plant-derived N varied between 89 and 102%. The results present evidence that the ability of the soil to retain plant-derived N is strong in comparison with the ability of the subsequent crop and different loss mechanisms to remove it.     

Effects of vesicular-arbuscular mycorrhiza on the availability of iron phosphates to plants

Summary The effect of inoculation with a mycorrhizal fungus on the growth of subterranean clover and of ryegrass was measured using three sources of phosphorus with different solubilities. These were (in order of decreasing solubility): potassium dihydrogen phosphate, colloidal iron phosphate and crystalline iron phosphate. Mycorrhizal infection increased growth more for subterranean clover than for ryegrass for all sources of phosphorus. For both species the greatest benefit from mycorrhizal inoculation was obtained with the least soluble source of iron phosphate. It is suggested that the mycorrhizas were able to explore the soil more thoroughly and hence were able to locate and use the point sources of phosphorus in the insoluble iron phosphates.     

A STUDY OF THE BACTERIA ASSOCIATED WITH THE ROOTS OF SUBTERRANEAN CLOVER AND WIMMERA RYE-GRASS

SUMMARY: A study of the bacteria from the surfaces of roots of subterranean clover ( T. subterraneum L.) and Wimmera rye-grass ( L. rigidum Gaud.) revealed that 21 genera were represented among the isolates from clover and 16 genera among those from rye-grass. Bacteria showing branched forms predominated and accounted for 63% of the 151 clover isolates and 78% of the 167 grass isolates. Most of these were identified as Arthrobacter , but from clover a significant proportion were Nocardia -like types. Members of the genera Mycoplana, Micromonospora, Mycobacterium , and Mycococcus were also identified among the branching forms.
Although the soil had been inoculated with effective rhizobia and the clover plants were effectively nodulated only one of the 318 isolates was capable of nodulating subterranean clover. The majority of the isolates were chromogenic and Gram-negative, produced acid from glucose and ammonia from peptone, were catalase-positive and grew best aerobically. Approximately half the isolates liquefied gelatin and produced hydrogen sulphide from peptone.     

Improving N2 fixation from the plant down: Compatibility of Trifolium subterraneum L. cultivars with soil rhizobia can influence symbiotic performance

Plant genotypes of Trifolium subterraneum L. (subterranean clover) were evaluated for differences in symbiotic N₂ fixation with soil rhizobia, with the long-term aim of using plant selection to overcome sub-optimal N₂ fixation associated with poorly effective soil rhizobia. Symbiotic performance (SP) was assessed for 49 genotypes of subterranean clover with each of four pure Rhizobium strains isolated from soil. Plants were grown in N free media in the greenhouse and their shoot dry weights measured and expressed as a percentage of dry weight with R. leguminosarm bv. trifolii WSM1325, the recommended commercial inoculant. Average SP with two Rhizobium strains (H and J) ranged from completely ineffective to 80% of potential for the subterranean clover genotypes. Two clover cultivars with high (cv. Campeda) and low (cv. Clare) SP values were investigated in more detail. Campeda typically fixed more N₂ than Clare when inoculated with 30 soil extracts (4.2 vs 2.4 mg N₂ fixed/shoot) and with 14 pure strains isolated from those soils (4.2 vs 2.2 mg N₂ fixed/shoot). The poor performance of Clare could be attributed to interruptions at multiple stages of the symbiotic association, from nodule initiation (less nodules), nodule development (small, white nodules), through to reduced nodule function (N₂ fixed/mg nodule) depending on the inoculation treatment. Through the careful use of subterranean clover genotypes by plant breeders it should be possible to make significant gains in the SP of future subterranean clover cultivars.     

Hydrogen and aluminium tolerance

Summary Reduced productivity due to soil acidity has been demonstrated with subterranean clover and wheat in many parts of Australia. Nodulation in clover appears to be more sensitive to low pH than growth of the host plant in the presence of adequate mineral nitrogen. Low pH is associated with aluminium toxicity in a number of species and nodulation in clover is more sensitive than growth of the host plant to Al. Decrease in soil pH is associated with significant increases in exchangeable Al. The breeding of Al tolerant wheats in Australia involves a backcross programme utilizing the transfer of tolerance from the Brazilian cultivar Carazinho.     

Seed-borne cucumber mosaic virus infection of subterranean clover in Western Australia

In Western Australia, infection with cucumber mosaic virus (CMV) was widespread in all three subspecies of subterranean clover (Trifolium subterraneum) growing in plots belonging to the Australian National Subterranean Clover Improvement Programme. Seed-borne CMV was detected in seed harvested in 1984–1986 of 18/25 cultivars from two collections of registered cultivars; seed transmission rates ranged up to 8.8%. Seed samples from CMV-inoculated plants of 11 cultivars transmitted the virus to 0.5–8.7% of seedlings. Seed transmission rates greater than 5% were obtained only with cvs Enfield, Green Range and Nangeela. CMV was not detected in seed harvested in 1975–1981 from one of the registered cultivar collections, in 17 commercial seed stocks from 1986 or in a survey of subterranean clover pastures.
Symptoms in subterranean clover naturally infected with CMV included mottle, leaflet downcurling and dwarfing but severity varied with cultivar and selection. CMV isolates from different sources varied in virulence when inoculated to subterranean clover; two (both from subterranean clover) were severe, two moderate and three (including one from subterranean clover) mild. In pot tests, CMV decreased herbage production and root growth (dry wts) of cv. Green Range by 49% and 59% respectively. In spaced-plants growing in plots, CMV decreased herbage production and root growth of cvs Green Range and Northam by 59–630 and seed production of cv. Green Range by 45%. In rows sown with infected seed, aphid spread increased infection levels to 75% in cv. Green Range and 44% in cv. Esperance and losses in herbage production of 42% and 29% respectively were recorded.
CMV isolated from subterranean clover included isolates from both serogroups.     

Competition between Subterranean Clover and Rygrass for uptake of 15N-labeled fertilizer

Grasses and legumes are often grown together for improving quality of forage and for better yield when soil N availability is limiting. One compatible mixture is Trifolium subterranium L., subterranean clover and Lolium multiflorum Lam, ryegrass.Experiments were conducted with plants grown in a glasshouse and plant growth chambers to determine the competitive ability of these plants for fertilizer N. Fertilizer N was enriched with ¹⁵N to measure the contribution of dinitrogen fixation and fertilizer N to the growth of clover. In pure stands, with increased fertilizer N, the legume took up similar quantities of mineral N as the grass to make up for the deficit due to less dinitrogen fixation but in mixed stands the grass by far outcompeted the legume. The growth of clover suffered due to lack of N both from less dinitrogen fixation and the inability to compete with the grass for mineral N. Increasing levels of fertilizer N reduced dinitrogen fixation by the clover. When growing with the clover the grass did not receive N from the clover. A laboratory experiment using ¹⁵N label on pure stands of the two species indicated that the grass had an inherent capability of absorbing almost twice the amount of mineral N as the legume under the same conditions even when root weight and volume was not larger for the grass. The results of this research provide insight into the often observed phenomenon that growth of clover is reduced when grown with grass in proportion to the amount of mineral N provided. This revised version was published online in June 2006 with corrections to the Cover Date.