Differential response of citrus rootstocks to aluminum levels in nutrient solutions: II. Plant mineral concentrations 2

The objective of this study was to determine relations between Al effects and mineral concentrations in citrus seedlings. Six‐month‐old seedlings of five citrus rootstocks were grown for 60 days in supernatant nutrient solutions of Al, P, and other nutrients. The solutions contained seven levels of Al ranging from 4 to 1655 μM. Al and similar P concentrations of 28 μM P. Aluminum concentrations in roots and shoots increased with increasing Al concentration in the nutrient solution. Aluminum concentrations in roots of Al‐tolerant rootstocks were higher than those of Al‐sensitive rootstocks. When Al concentrations in nutrient solution increased from 4 to 178 μM, the K, Mg, and P concentrations in roots and the K and P levels in shoots increased. Conversely, Ca, Zn, Cu, Mn, and Fe in the roots and Ca, Mg, Cu, and Fe in the shoots decreased. The more tolerant rootstocks contained higher Fe concentrations in their roots than did the less tolerant ones when Al concentrations in solution were lower than 308 μM. Concentrations of other elements (Ca, K, P, Mg, Zn, and Mn) in roots or shoots exhibited no apparent relationship to the Al tolerance for root or shoot growth of the rootstocks. Calcium, K, Zn, Mn, and Fe concentrations in roots and Mg and K concentrations in shoots of all five rootstocks seedlings had significant negative correlations with Al concentrations in corresponding roots or shoots.     

Growth and Ca,Mg, K,and P uptake by triticale,wheat, and rye at four al levels

This study was conducted to determine relationships between Al toxicity and mineral uptake of triticale (X Triticosecale, Wittmack), wheat (Triticum aestivum L.), and rye (Secale cereale L.). Two culti‐vars of each species were grown in 1/5‐strength Steinberg solution with 0, 3, 6, or 12 ppm Al added. The solutions were adjusted to pH 4.8 at transplanting and were not adjusted thereafter. The plants were grown in a growth chamber for 19 days before harvesting to determine nutrient solution pH, dry weights, and Al, Ca, Mg, K, and P levels in plants. Increasing Al concentration reduced the final pH of solutions. The addition of 12 ppm Al severely reduced the growth and increased Al concentration of plant tops. The Al levels in roots generally increased with increments of added Al up to 6 ppm. Increasing Al decreased the uptake of Ca, Mg, and P by plant tops more than that of K. Regression analyses indicated that Al toxicity was associated with increasing K/Ca + Mg equivalent ratios and decreasing P concentration in plant tops. Differences between species were: higher Al concentration in rye than wheat with 6 and 12 ppm Al, higher translocation of Ca from roots to tops in wheat than in rye and Mg in triticale and wheat than rye; K/Ca + Mg equivalent ratios associated with 50% reduction in top growth followed the order: triticales > tolerant wheat > sensitive wheat > rye. Differences in mineral uptake associated with Al toxicity in wheat were more indicative of differential Al sensitivity in wheat than in triticale and rye which have higher internal Al tolerance.     

Effect of Glomus etunicatum inoculation on aluminum,phosphorus, calcium,and magnesium uptake of two barley genotypes with different aluminum tolerance

Abstract

This study was conducted to evaluate the effect of vesicular‐arbuscular mycorrhizal (VAM) fungus Glomus etunicatum on growth, absorption, and distribution of calcium (Ca), magnesium (Mg), phosphorus (P), and aluminum (Al) in one Al‐tolerant and one Al‐sensitive barley cultivar. The plants were grown in sand daily irrigated with nutrient solution containing 0 or 600 μM Al at pH 4.8. Significant interaction (P=0.05) among variety, mycorrhiza, and aluminum (VxMxAl) were noted for both shoot and root dry matter (DM); shoot concentration and content of Al, P, Ca, and Mg; root concentration of Al, P, and Mg; and root content of Al, P, Ca, and Mg. With VAM inoculation: i) root colonization degree was about 50% in all treatment, ii) shoot DM yield increased between 30 and 70%, iii) Al concentration and content decrease down to a half both in shoots and roots of sensitive barley, iv) Ca concentration in shoots of sensitive barley showed a high increase at 600 μM Al, and v) P concentration and content in shoots of both varieties increased significantly.     

Differential response of two taro cultivars to aluminum: II. Plant mineral concentrations

Abstract

One proposed mechanism of aluminum (Al)‐tolerance involves the ability of plants to maintain uptake of essential mineral elements in the presence of Al. To examine this hypothesis, taro [Colocasia esculenta (L.) Schott] cultivars (cv.) Lehua maoli and Bun long were grown in hydroponic solution at six initial Al levels (0, 110, 220, 440, 890, and 1330 μM Al), and plant mineral concentrations were determined after 27 days. Increasing Al levels significantly increased Al concentrations in taro leaf blades, petioles, and roots. This increase in Al concentrations in the leaf blades as solution Al levels increased was greater for Al‐sensitive cv. Bun long compared to cv. Lehua maoli, resulting in significant interaction between Al and cultivar effects. However, no significant cultivar differences were found for Al concentrations in the petioles or roots. Increasing Al levels in solution significantly depressed concentrations of calcium (Ca), magnesium (Mg), manganese (Mn), and iron (Fe) in taro leaf blades, and significantly depressed concentrations of Ca, Mg, copper (Cu), and zinc (Zn) in taro roots. Aluminum‐induced Ca deficiency appeared to be one possible mechanism of Al phototoxicity in taro, becvasue Ca concentrations in the leaf blades and roots at the higher Al levels were within the critical deficiency range reported for taro. Significant cultivar differences were found, in which Al‐tolerant cv. Lehua maoli had significantly greater Ca and Cu concentrations in the roots, and significantly greater potassium (K) concentrations in the leaf blades across all Al levels. Our results show that Al‐tolerance in taro cultivars is associated with the ability to maintain uptake of essential mineral nutrients, particularly Ca and K, in the presence of Al.     

Differences in aluminium tolerance of ryegrass and white clover plants grown from seed or propagated vegetatively

The effect of aluminium (Al) on the relative yield of plants grown from seeds of ryegrass (Lolium perenne L.) or white clover (Trifolium repens L.) and either tillers (ryegrass) or stolon tip cuttings (white clover) were investigated using a low ionic strength (2.7 x 10^‐3 M) solution culture technique. In ryegrass, plants grown from tillers had higher relative yields than plants grown from seedlings in the tops when solution Al concentrations were greater than 16 μM and in the roots when solution Al concentrations were greater than 7 μM. In white clover, relative yields in the tops and roots plants were higher in plants grown from stolon tip cuttings than from seedlings when solution Al concentrations were greater than 10 μM. There were no significant cultivar effects. The results indicate that plants used in Al‐tolerance experiments can be grown from seed or vegetatively propagated, provided solution Al rates are adjusted to reflect differences in Al tolerance.     

Aluminum-induced growth stimulation in relation to calcium,magnesium, and silicate nutrition in Melastoma malabathricum L.

Effects of Al, Ca, Mg, and Si on the growth and mineral accumulation of M. malabathricum (Melastoma malabathricum L.), which is an Al accumulator plant, were investigated using the water culture method. Rice (Oryza sativa L.) and barley (Hordeum vulgare L.) were used as control plants. After Al application, growth was inhibited in barley, but stimulated in M. malabathricum and rice. The growth of M. malabathricum was not reduced by very low Ca and Mg concentrations (0.1 mm Ca and 0.05 mm Mg). However, it was depressed in the absence of Ca. Ca and Mg contents somewhat decreased by Al application, which was most evident in young leaves and roots. M. malabathricum accumulated more than 10,000 mg kg^-1 Al in mature leaves, and more than 7,000 mg kg^-1 even in the youngest leaf. Al content in leaves of M. malabathricum did not decrease by the Ca or Mg application, but slightly decreased by in the absence of Ca.

Although Si is a strong ligand of Al in solution, in M. malabathricum, Si application hardly affected the growth, Al accumulation and nutrient uptake.     

Role of root cation‐exchange capacity in differential aluminum tolerance of lotus species

Three experiments were conducted in which roots of two species of Lotus were immersed for up to 40 min in complete nutrient solutions containing 6, 15 or 25 μM Al. The two species tested were L. pedunculatus cv. Grasslands Maku (Al‐tolerant) and L. corniculatus cv. Maitland (Al‐sensitive). There was an initial rapid (< 5 min) decrease in solution Al at 25 μM Al. The effect was less marked with solution Al ≤ 15 μM. The decrease in solution Al was greater in the Al‐sensitive Maitland than in the Al‐tolerant Grasslands Maku, particularly when expressed on the basis of root fresh mass and root length. Root cation‐exchange capacity (CEC) was lower in Grasslands Maku than in Maitland, viz. 23.9 vs 36.5 mmol kg^‐1 dry mass. Maitland roots removed more Al from solution than did those of Maku on the basis of total exchange capacity.

We propose a mechanism of Al tolerance on the basis of the results of this study and of other published information, viz. that differential Al tolerance results from differences in root CEC. Aluminum‐tolerant genotypes have roots with low CEC, and high Al activities (> 20 μM in the case of Grasslands Maku) are required to precipitate the relatively highly methylated pectins associated with low CEC. In contrast, relatively low activities of Al would precipitate the pectins in plants with roots of high CEC. This would decrease the protective capacity of the pectins, enabling the toxic, monomeric Al ions to come in contact with a number of Al‐sensitive compounds or processes in the cell wall, plasmalemma, or cell cytoplasm.     

Mineral element concentration of two barley cultivars in relation to water deficit and aluminum toxicity

The separate and combined effects of water and Al stress on concentrations of P, K, Ca, Mg, Fe, Mn, Zn, Cu, B, Al, Sr, and Ba were determined in tops of ‘Dayton’ (Al‐tolerant) and ‘Kearney’ (Al‐sensitive) barley (Hordeum vulgäre L.) grown in an acid, Al‐toxic, Tatum subsoil (clayey, mixed, thermic, Typic Hapludult). Plants were grown 4 weeks in a plant growth chamber at high (pH 4.7) or low (pH 6.6) Al stress. During the last 2 weeks they were also subjected to low (‐20 to ‐40 kPa), moderate (‐40 to ‐60 kPa), or high (‐60 to ‐80 kPa) water stress. In general, Al stress had a greater overall effect on mineral element concentration of tops than water stress. Aluminum stress significantly decreased concentrations of P, Ca, and Mg and increased concentrations of Zn, Sr, and Ba, irrespective of the cultivar or water stress treatment. Cultivar differences in Mn concentration were observed with Al stress under all water stress conditions. In each case, Mn concentration was lower in ‘Kearney’ than in ‘Dayton’. Potassium, Ca, and Mg were lower in ‘Kearney’ than in ‘Dayton’ only at low and moderate water stress, under low Al stress, ‘Kearney’ had significantly higher concentrations of K and Ca than did ‘Dayton’ under all water stress conditions. The effects of water stress on mineral element concentration varied greatly with cultivar, Al stress treatment, and severity of water stress. Under high Al stress, increasing drought conditions from low water stress (‐20 to ‐40 kPa) to high water stress (‐60 to ‐80 kPa) significantly increased the concentrations of Ca, K, Zn, Sr, and Ba in Al‐sensitive ‘Kearney’ and reduced the concentrations of Zn, Sr, and Ba in Al‐tolerant ‘Dayton'; P and Mg concentration were unaffected by water stress. In contrast, under low Al stress, a corresponding increase in water stress significantly increased the concentrations of Ca and reduced that of P in ‘Kearney’ and increased Ca and B concentration in ‘Dayton'; Mg concentrations were unaffected in either cultivar. Thus, it appears that Al stress and water stress had opposite effects on Ca accumulation in barley tissue.     

Effect of manganese on concentrations of Zn,K, Ca and Mg in two bush bean cultivars grown in solution culture

Two bush bean cultivars [Phaseolus vulgaris L. cv. ‘Wonder Crop 2’ (WC‐2) and ‘Green Lord’ (GL)], differing in Mn toxicity, were grown in a growth chamber for 12 days in Hoagland No. 2 nutrient solution containing 0.05 to 1 ppm Mn as MnCl₂4H₂O with 1 ppm Fe as Fe‐EDTA, at an initial pH 5.00. Concentrations of Zn, K, Ca and Mg in the tissues of two bush bean cultivars were examined in relation to Mn toxicity.

The concentration of Zn in the leaves of Mn‐sensitive WC‐2 increased significantly with increasing Mn concentration in the solution, but such levels were not toxic to the plants.

The percent distribution of Zn and K in Mn‐sensitive WC‐2 plants (% of total uptake) significantly increased in the tops and decreased in the roots with increasing Mn concentration in the nutrient solution; however, Mn treatment had no effect on distribution of either Ca or Mg in WC‐2. External Mn concentration had little or no effect on the K, Ca, or Mg concentration in the tops of Mn‐tolerant GL.     

Effects of low aluminum activity in nutrient solutions on the organic acid concentrations in maize plants

The effects of low aluminum (Al) activity in nutrient solution on the concentrations of organic acids in two cultivars of maize (Zea mays L.), HS7777 Al‐sensitive and C525‐M Al‐tolerant, were studied. Aluminum stress increased total organic acid concentration in the roots and in the shoots for both cultivars. The relative increase of t‐aconitic, citric, formic, malic, and quinic acids was higher in the roots than in the shoots for both cultivars. The concentrations of c‐aconitic, isocitric, malonic, oxalic, and succinic organic acids were reduced by Al stress, principally for C525‐M. There were no consistent differences in organic acid concentrations between the cultivars to discriminate Al tolerance. The Al tolerance for C525‐M may be justified by lower Al concentrations in the root tips where cellular division takes place and/or by higher excretion of organic acids from roots to the rhizosphere for detoxification of Al by chelation.     

Effects of low aluminium levels on growth and nutrient relations in three rice cultivars with different tolerances to aluminium

Reports on varietal diversity of upland rice in relation to relatively low aluminium (Al) levels are limited. Therefore, effects were examined of 35, 70, and 140 μM Al on plant growth and uptake of macro‐ and micro‐nutrients (K, P, Ca, Mg, Fe, Zn, Cu, and Mn) and their distribution in three upland rice (Oryza saliva L.) cultivars (BG35, DA14, and IR45) with different Al sensitivity. After an initial growth period of 5 days without Al, the plants were grown for 21 days in nutrient solutions containing Al at pH 4.1. Cultivar BG35 showed the highest and IR45 the lowest tolerance to Al when fresh weights of shoots or roots were considered. Except for IR45 at 140 μM Al, total dry weight was unaffected by Al, and the cultivars could not be clearly distinguished with respect to Al tolerance. Net Al uptake rate was higher in Al tolerant BG35 than in DA14 or IR45. Conversely, in IR45 the absorbed Al was rapidly transported to the shoots and accumulated there. In BG35, net P and Ca uptake rates in Al‐treated plants were high enough to maintain the P and Ca status of the shoots at all Al levels. Irrespective of Al sensitivity, there was a general depression of internal Mg concentration in Al‐reated plants. The Fe, Zn, Cu, and Mn concentrations of the plants were not negatively affected by Al in any of the cultivars.     

Effect of aluminium on onions,asparagus and squash

The effect of Al on the growth and chemical composition of onions (Allium cepa L. cv Pukekohe longkeeper), asparagus (Asparagus officinalis L. cv Lucullus) and squash (Cucurbita maxima L. cv Delica) was investigated using a low ionic strength solution‐culture technique. Onions and asparagus were sensitive to Al while squash was relatively tolerant. The dry matter yields of onion, asparagus and squash tops were reduced 50% by 5, 3, and 26 μM Al in the solution, respectively. Similarly, root dry matter yields were reduced 50% by 1, 5, and 26 μM Al, respectivley. Aluminium reduced the concentration of macro nutrients in both asparagus and onion tops but had no consistent effect on squash tops. Plant analysis and visible symptoms indicated that Al affected magnesium nutrition more than that of any other nutrient.

The roots of squash showed an ability to recover from Al stress by modifying their morphology. However, this effect was insufficient to totally alleviate Al toxicity. Squash also became more sensitive to Al as the growing period progressed. These observations suggest that care is needed in using short term experiments to assess Al tolerance for some species.     

Varietal diversity of upland rice in sensitivity to aluminium

A rapid and simple nutrient addition technique was used for evaluating Al tolerance of six local upland rice (Oryza sativa L.) cultivars (BG35, BR21, DA25, DA26, DA14, and DA22) from Bangladesh and three IRRI rice, IR46, IR97, and IR45, cultivars from the Philippines. The plants were grown for 21 days with Al (0 μM, 140 μM, 280 μM or 560 μM) at pH 4.1. The roots were more affected by Al than the shoots. In rating cultivars for Al sensitivity, relative shoot weight (RSW) was found to be the best parameter due to the severe damage of the roots, irrespective of Al sensitivity. The cultivars were rated as Al tolerant (BG35, BR21, DA25, and DA26), mid‐tolerant (DA14, DA22, and IR46) and sensitive (IR97 and IR45) . More Al was retained in the roots of tolerant cultivars than in the mid‐tolerant or sensitive cultivars. In shoots, the Al concentration of tolerant cultivars was less than in the mid‐tolerant or in the sensitive cultivars and the inhibition of growth was proportional to Al concentration irrespective of Al tolerance. Therefore, the variation among cultivars in Al sensitivity could be related to the capacity of roots to retain Al from transport to the shoots.     

Variation in aluminum tolerance among and within lotus lines

Previous studies with one cultivar of each of two Lotus species showed that L. pedunculatus cv. Grasslands Maku is Al‐tolerant while L. corniculatus cv. Maitland is Al‐sensitive. However, little information exists on the relative Al‐tolerance of other lines in these species or on the variability within lines in response to Al. Two solution culture experiments were conducted to study these aspects. The Al tolerance of Grasslands Maku was confirmed, plant dry mass increasing by > 40 % with 10 to 30 μM monomeric Al in solution. In contrast, the yield of Maitland was decreased by > 80 % at 30 μM Al, as was that of the other two L. corniculatus lines tested, AT‐I and AT‐U. The F1 of a cross between a L. pedunculatus line and a L. corniculatus line (G4712) was intermediate with respect to Al tolerance. Also, there was a wide range of response to Al in the F2, which appeared to have similar morphological characteristics to Maitland. Thus, there would appear to be considerable benefit in selecting for Al tolerance and good agronomic characters in this interspecific cross.     

The influence of acid soil factors on the growth of snapbeans in major appalachian soils

Abstract

Acid soil limitations to plant growth were assessed In 55 horizons of 14 major Appalachian hill land soils. Aluminum sensitive “Romano” and Al‐tolerant “Dade” snapbeans (Phaseolus vulgaris L.) were grown for 5 weeks in limed and unlimed treatments of the 55 horizons. Shoot and root growth was depressed >20% in unlimed relative to limed treatments in approximately 2/3 of the horizons. Dade snapbeans were generally more tolerant of the acid soil conditions and had higher Ca concentrations in the shoots than Romano snapbeans. However, the sensitive‐tolerant snapbean pair could not consistently be used to identify horizons with soil Al problems. Growth of both snapbeans was generally best in A horizons and worst in E horizons. The E horizons in this study were characterized by low Ca saturation (exchangeable Ca x 100/cation exchange capacity) and high Al saturation (exchangeable Al x 100/cation exchange capacity). Exchangeable Ca, soil Ca saturation and total soil solution Ca were positively correlated (p<0.01) with snapbean root and shoot growth. Soil Al saturation, total soil solution Al and soil solution Al reacting in 15 seconds with 8‐hydroxyquinoline were negatively correlated (p<0.01) with growth. The ratio of Ca/Al in soil solution was more closely related to snapbean growth than the soil solution concentration of any individual element. Soil and soil solution Mn were, in general, not significantly correlated with snapbean growth. Many of the horizons in this study had both Al toxicity and Ca deficiency problems and interaction between Ca and Al affected both snapbean growth and Ca uptake. These findings confirm the importance of considering Ca as well as Al when investigating Al phytotoxicity.     

Short‐term effects of pH and aluminium on mineral nutrition in maize varieties differing in proton and aluminium tolerance

In short‐term (24 h) nutrient solution experiments, the influence of different proton (pH 6.0 and pH 4.3) and aluminium (Al) (0, 20, and 50 μM) concentrations on root and coleoptile elongation, dry weight, and the uptake of selected mineral nutrients was studied in maize (Zea mays L.) varieties that differ in acid soil tolerance under field conditions. The acid‐soil‐tolerant maize varieties, Adour 250 and C525M, proved to be hydrogen (H⁺) ion sensitive, but Al tolerant, while the acid soil tolerant variety BR201F was H⁺ tolerant but Al sensitive. The acid soil sensitive variety HS 7777 was affected by both H⁺ and Al toxicity. The proton‐induced inhibition of root elongation was closely related to the proton‐induced decrease of the specific absorption rates (SAR) of boron (B), iron (Fe), magnesium (Mg), calcium (Ca), and phosphorus (P). In contrast, only the specific absorption rate of B (SARB) was significantly correlated to the Al‐induced inhibition of root elongation. It is concluded, that alterations of nutrient uptake may play an important role in H⁺ toxicity, while at least after short‐term exposure to Al, alterations of Ca, Fe, Mg, or P uptake do not seem to be responsible for Al‐induced inhibition of root elongation. Further attention deserves the Al‐B interaction, moreover taking into account that a highly significant correlation between Al‐induced increase of callose concentration in root tips and Al‐induced decrease of SAR_B could be established.     

Aluminium toxicity with sorghum genotypes in nutrient solutions and its amelioration by magnesium

Effects of Al toxicity and interaction of Al and Mg on growth of twelve sorghum (Sorghum bicolor (L.) Moench) genotypes have been studied in nutrient solutions (pH 4.2). Aluminium at 30 μM decreased biomass (dry matter yield) of the individual genotypes by factors between 1.27 and 7.36, with identical sensitivity grouping of genotypes as obtained in an earlier pot experiment with an acid soil. Resembling acid-soil stress, Al toxicity was simultaneously expressed in two independent ways, i.e. impairment of root development and induced Mg deficiency. The effect of Al on total dry matter production of the genotypes was correlated more closely with changes in specific root length (m g⁻¹ dry root) than with changes in internal Mg status. Increased Mg concentrations in the solutions (2.5 and 7.5 instead of 0.25 mM) not only decreased Al-induced Mg deficiency but also reduced the concentrations of Al in/on the roots and its damaging effect on root development. Therefore, the sorghum genotypes were less sensitive to Al at the higher Mg levels. At a high Mg concentration in the solution (7.5 mM) dry matter yield of two genotypes was even stimulated by Al.     

Effects of alumumium on nutrient composition and yield of oats

Spotty chlorosis appeared on the young leaves of Al toxic plants and was severe or high Al level. Stems of the Al‐treated plants were purple While the roots were short, thick and brown in colour with little branching.

The dry matter yield of tops and roots decreased with increased Al level. The concentration of P was greater in roots of Al toxic plants than in the control plants but a converse affect was recorded in tops.

Increased rates of Al caused a decrease in the concentrations of P,K,Ca, M_c and Mn in plant tops, while Al concentration increased both in tops and roots. Iron content in tops affected with increased Al and in root a regular increase was observed. The results indicate that Al toxicity depressed the growth of oats, resulted in abnormal root development with tittle branching and affecting the utilization of several essential plant nutrients by oat plant.     

Growth and nutrient uptake of peach seedlings with varying magnesium concentrations at low pH 1

Soils of the peach growing region of the Southeastern Coastal Plain are highly leached and excessively acid, with inherently low concentrations of subsoil magnesium (Mg). A greenhouse experiment was conducted to determine the effects of varying Mg concentrations at low pH on growth and Mg uptake of three peach seedling cultivars commonly used as rootstock in the region. Seedlings of ‘Lovell’, ‘Elberta’, [Prunus persica (L.) Batsch] and ‘Nemaguard’ [Prunus persica (L.) Batsch X Prunus davidiana Carriere] were grown for 36 days in nutrient solution containing 9, 21, 42, 84, 167, 333, and 667 μM Mg. Magnesium concentration in solution did not increase lateral length, number of laterals, trunk cross‐sectional area, or root volume. All terminal growth responses were cultivar related. Magnesium concentration in the leaves, stems, and roots were increased either by quadratic or cubic relationship with solution Mg concentration while Mg uptake rate was increased linearly with solution Mg concentration with all three seedling cultivar. Uptake rates of calcium, manganese, and zinc, and tissue concentrations of phosphorus, manganese, and zinc decreased with increasing Mg concentrations in nutrient solution. Predicted Mg uptake rates by‐regression analysis revealed a cubic uptake isotherm for Nemaguard and a quadratic isotherm for Elberta. Predicted tissue Mg concentration followed similar patterns of accumulation for leaves and stems, but root Mg concentration followed a cubic uptake isotherm for all three seedlings. The linear Mg uptake at low pH may be an important physiological characteristic that enables Lovell seedlings to outperform either Elberta or Nemaguard when used as a rootstock in the southeastern soils.