Soil Chemical Analysis

Toxic metal pollution of waters and soils is a major environmental problem, and most conventional remediation approaches do not provide acceptable solutions. The use of specially selected and engineered metal-accumulating plants for environmental clean-up is an emerging technology called phytoremediation. Three subsets of this technology are applicable to toxic metal remediation: (1) Phytoextraction--the use of metal-accumulating plants to remove toxic metals from soil; (2) Rhizofiltration--the use of plant roots to remove toxic metals from polluted waters; and (3) Phytostabilization--the use of plants to eliminate the bioavailability of toxic metals in soils. Biological mechanisms of toxic metal uptake, translocation and resistance as well as strategies for improving phytoremediation are also discussed.

Phytoremediation: A Novel Strategy for the Removal of Toxic Metals from the Environment Using Plants

Plant developmental processes are controlled by internal signals that depend on the adequate supply of mineral nutrients by soil to roots. Thus, the availability of nutrient elements can be a major constraint to plant growth in many environments of the world, especially the tropics where soils are extremely low in nutrients. Plants take up most mineral nutrients through the rhizosphere where micro-organisms interact with plant products in root exudates. Plant root exudates consist of a complex mixture of organic acid anions, phytosiderophores, sugars, vitamins, amino acids, purines, nucleosides, inorganic ions (e.g. HCO3−, OH−, H+), gaseous molecules (CO2, H2), enzymes and root border cells which have major direct or indirect effects on the acquisition of mineral nutrients required for plant growth. Phenolics and aldonic acids exuded directly by roots of N2-fixing legumes serve as major signals to Rhizobiaceae bacteria which form root nodules where N2 is reduced to ammonia. Some of the same compounds affect development of mycorrhizal fungi that are crucial for phosphate uptake. Plants growing in low-nutrient environments also employ root exudates in ways other than as symbiotic signals to soil microbes involved in nutrient procurement. Extracellular enzymes release P from organic compounds, and several types of molecules increase iron availability through chelation. Organic acids from root exudates can solubilize unavailable soil Ca, Fe and Al phosphates. Plants growing on nitrate generally maintain electronic neutrality by releasing an excess of anions, including hydroxyl ions. Legumes, which can grow well without nitrate through the benefits of N2 reduction in the root nodules, must release a net excess of protons. These protons can markedly lower rhizosphere pH and decrease the availability of some mineral nutrients as well as the effective functioning of some soil bacteria, such as the rhizobial bacteria themselves. Thus, environments which are naturally very acidic can pose a challenge to nutrient acquisition by plant roots, and threaten the survival of many beneficial microbes including the roots themselves. A few plants such as Rooibos tea (Aspalathus linearis L.) actively modify their rhizosphere pH by extruding OH− and HCO3− to facilitate growth in low pH soils (pH 3 – 5). Our current understanding of how plants use root exudates to modify rhizosphere pH and the potential benefits associated with such processes are assessed in this review.

Root exudates as mediators of mineral acquisition in low-nutrient environments

High variability in plant-growth response to the presence of different mycorrhizal fungi can be a major determinant of local plant species diversity. Multiple species of arbuscular mycorrhizal fungi can coexist in terrestrial ecosystems, and co-occurring plants can differ in their response to colonization by these different fungi. However, the range of mycorrhizal plant-growth responses that can occur within communities has not been determined. In the present study, I crossed a large number of plant and fungal species that co-occur to determine the range of responses that can exist within an ecosystem. I also crossed exotic fungal isolates vs. local plant isolates and local fungal isolates vs. exotic plant isolates to determine whether the range of plant growth responses differs when using foreign genotypes. The data indicate that plant growth responses to mycorrhizal inoculation within an ecosystem can range from highly parasitic to highly mutualistic. In this study, the direction and magnitude of the response depended on the combination of plant and fungal species. No plant did best with the same fungal isolate. The range of responses was greatest when using local plants and fungi. Whereas parasitic and mutualistic responses were also detected when using foreign plant or fungal genotypes, the range of responses was significantly reduced, as was the relative frequency of positive responses. Overall, this study suggests that, within ecosystems, arbuscular mycorrhizal fungi can function along a continuum from parasitism to mutualism, and that extreme responses are more common when using locally adapted plants and fungi. This high variation in plant growth response may be a large contributor to plant species coexistence and the structure of plant communities.

Variation in plant response to native and exotic arbuscular mycorrhizal fungi

Understanding how terrestrial ecosystems function requires a combined aboveground–belowground approach, because of the importance of feedbacks that occur between herbivores, producers, and the decomposer subsystem. In this paper, we identify several mechanisms by which herbivores can indirectly affect decomposer organisms and soil processes through altering the quantity and quality of resources entering the soil. We show that these mechanisms are broadly similar in nature for both foliar and root herbivory, regardless of whether they operate in the short term as a result of physiological responses of individual plants to herbivore attack or long-term following alteration of plant community structure by herbivores and subsequent changes in the quality of litter inputs to soil. We propose that a variety of possible mechanisms is responsible for the idiosyncratic nature of herbivore effects on soil biota and ecosystem function; positive, negative, or neutral effects of herbivory are possible depending upon the balance of these different mechanisms. However, we predict that positive effects of herbivory on soil biota and soil processes are most common in ecosystems of high soil fertility and high consumption rates, whereas negative effects are most common in unproductive ecosystems with low consumption rates. The significance of multiple-species herbivore communities is also emphasized, and we propose that if resource use complementarity among herbivore species or functional groups leads to greater total consumption of phytomass, and thus greater net herbivory, then both positive and negative consequences of increasing herbivore diversity for belowground properties and processes are theoretically possible. Research priorities are highlighted and include a need for comparative studies of herbivore impacts on above- and belowground processes across ecosystems of varying productivity, as well as a need for experimental testing of the influence of antiherbivore defense compounds on complex multitrophic interactions in the rhizosphere and the significance of multiple herbivore species communities on these plant–soil interactions.

Herbivore-mediated linkages between aboveground and belowground communities

Iron deficiency chlorosis (FeDC) is a common disorder for sorghum [Sorghum bicolor (L.) Moench] grown on alkaline calcareous soils. Four sorghum genotypes were grown in growth chambers on a low Fe (1.3 μg/g DTPA-extractable), alkaline (pH 8.0), calcareous (3.87% CaCO3 equivalent) Aridic Haplustoll to determine effects of different soil temperatures (12, 17, 22 and 27°C at a constant 27°C air temperature) on various root and shoot growth traits and development of FeDC. As soil temperature increased, leaf chlorosis became more severe, and shoot and root dry weights, root lengths, and leaf areas increased markedly. Shoot/root ratios, shoot weight/root length, leaf area /shoot weight and leaf area /root weight and root length also increased while root length/root weight decreased as soil temperature increased. Severe FeDC developed in all genotypes even though genotypes had previously shown different degrees of resistance to FeDC. Genotypes differed in most growth traits, especially dry matter yields, root lengths, and leaf areas, but most traits did not appear to be related to genotype resistance to FeDC. The most FeDC resistant genotype had the slowest growth rate and this may be a mechanism for its greater resistance to FeDC.

Effects of soil temperature on root and shoot growth traits and iron deficiency chlorosis in sorghum genotypes grown on a low iron calcareous soil

Sorghum [Sorghum bicolor (L.) Moench] plants were grown in growth chambers at 20, 25 and 30°C in a low P Typic Argiudoll (3.65 µg P g−1 soil, pH 8.3) inoculated with Glomus fasciculatum, Glomus intraradices, and Glomus macrocarpum to determine effects of vesicular-arbuscular mycorrhizal fungi (VAMF) species on plant growth and mineral nutrient uptake. Sorghum root colonization by VAMF and plant responses to Glomus species were temperature dependent. G. macrocarpum colonized sorghum roots best and enhanced plant growth and mineral uptake considerably more than the other VAMF species, especially at 30°C. G. fasciculatum enhanced shoot growth at 20 and 25°C, and mineral uptake only at 20°C. G. intraradices depressed shoot growth and mineral uptake at 30°C. G. macrocarpum enhanced shoot P, K, and Zn at all temperatures, and Fe at 25 and 30°C above that which could be accounted for by increased biomass. Sorghum plant growth responses to colonization by VAMF species may need to be evaluated at different temperatures to optimize beneficial effects.

Effects of species of VA-mycorrhizal fungi on growth and mineral uptake of sorghum at different temperatures

Sorghum [Sorghum bicolor (L.) Moench] was grown in a greenhouse in a low P (3.6 mg kg−1) soil (Typic Argiudolls) inoculated with the vesicular-arbuscular mycorrhizal fungi (VMAF) Glomus fasciculatum and P added at 0, 12.5, 25.0, and 37.5 mg kg−1 soil to determine the effects of VAMF-root associations on plant growth, benefit and cost analysis, and P efficiency (dry matter produced/unit P absorbed). Root colonization with V A M F and shoot growth enhancements decreased with increased soil P applications. Mycorrhizal plants were less P efficient than nonmycorrhizal plants. Shoot dry matter differences between mycorrhizal and nonmycorrhizal plants were considered the benefit derived by plants from VAMF-root associations. Shoot dry matter differences between mycorrhizal and nonmycorrhizal plants with similar P concentrations were considered the costs paid by plants for VAMF-root associations. Values of benefit and cost analysis for VAMF-root associations were highest when soil P was lowest and decreased with increasing P applications. Genotypic differences for calculated costs were pronounced, but not benefits. Benefit and cost analysis may be helpful to evaluate host plant genotypes and VAMF species to optimize efficiencies of VAMF symbiosis in different soil environments.

Benefit and cost analysis and phosphorus efficiency of VA mycorrhizal fungi colonizations with sorghum (Sorghum bicolor) genotypes grown at varied phosphorus levels

The role of Cu in promoting the reproductive phase of growth was examined using Thatcher spring wheat (Triticum aestivum L.). Plants were grown in limed Bladen soil (pH 5.3) in a controlled growth room. The plants were first harvested when five leaves had developed (day 23) and some top leaves of Cu-deficient (-Cu) plants had rolled (withertip) indicating a Cu deficiency. The leaves were shorter on Cu than Cu-sufficient (+Cu) plants, and fewer leaves developed. As they entered the reproductive growth phase, +Cu plants accumulated reducing sugars and reduced 2, 3, 5- triphenyltetrazolium chloride (TTC) in their stems, while Cu plants had lower concentration of the reducing sugars and very little TTC was reduced indicating a reduction in energy and in reducing capacity. Thin-layer-chromatography showed that top leaves of Cu plants contained higher concentrations of aspartic acid, alanine, and serine; and less aminobutyric acid than +Cu plants. Nitrate, P, and K concentrations were higher, and Ca and Cu were lower in Cu than in +Cu plants.

Copper as essential to wheat reproduction

Genetic manipulation of crops to tolerate mineral stresses is a practical approach to improve productivity of tropical acid soils. Both acid soil tolerant (AS-T) and susceptible (AS-S) sorghum [Sorghum bicolor (L.) Moench] genotypes were grown in the field on an acid ultisol at Quilichao, Colombia, South America at 60% (60-Al) and 40% (40-Al) Al saturation to evaluate plants for growth and yield traits.

R. B. Clark

Papers

Effects of soil temperature on root and shoot growth traits and iron deficiency chlorosis in sorghum genotypes grown on a low iron calcareous soil

Effects of species of VA-mycorrhizal fungi on growth and mineral uptake of sorghum at different temperatures

Benefit and cost analysis and phosphorus efficiency of VA mycorrhizal fungi colonizations with sorghum (Sorghum bicolor) genotypes grown at varied phosphorus levels

Copper as essential to wheat reproduction

Growth and yield traits of sorghum grown on acid soil at varied aluminum saturations