Genotypic Differences in Aluminum Response of Plants

Comparative studies of aluminum effects in 22 species in seven plant families have established that some species or genotypes within species can resist aluminum toxicity (82). Foy (165) proposed 'tailoring the plant to fit the soil; in other words, he suggested that it was more economical to develop mineral-stress-resistant plants than to correct the soil for nutrient deficiencies or toxicities. This statement is particularly true for acid subsoils, where it is not economically feasible to lime at such depths, or for developing countries, where farmers cannot afford the high-input costs of lime.

Screening Tests

Screening for genotypic differences in response to aluminum toxicity can be conducted in pots or in fields with aluminum-toxic soil. A more rapid screening test for differences in aluminum tolerance among species or genotypes within species utilizes the aluminum-induced inhibition of root elongation as a measure of aluminum sensitivity (166). These tests are conducted with varying levels of aluminum in solution at an acid pH (≤4.5) to maintain a high activity of Al3+, the phytotoxic ion. Some researchers have found a poor correlation between plant responses in soil with those in nutrient solution (167). Others have found a good correlation (168-171).

Hematoxylin stains extracellular aluminum phosphate compounds that result from aluminum damage to root cells (172). Another quick screening test is to stain roots grown in an aluminumcontaining solution with hematoxylin and to assess the intensity of staining (173). With wheat, Scott et al. (174) found a good agreement between root elongation results and those using hematoxylin. However, Bennet (175) warned that many aspects of hematoxylin staining are not well understood and that aluminum-treated roots do not always respond to hematoxylin even when symptoms of aluminum toxicity occurred. Further, sometimes roots will stain in the absence of aluminum (175).

Moore et al. (176) proposed that recovery of root elongation after 48 h of exposure to aluminum is a better measure of irreversible damage to the root apical meristem. Hecht-Buchholz (177) reported that aluminum toxicity in barley caused stunted roots, destruction of root cap cells, swelling, and destruction of both root epidermal and cortical cells. She found large differences between cultivars and proposed that aluminum resistance could be attributed to greater resistance of the root meristem of the aluminum-tolerant genotype to irreversible destruction. Lazof and Holland (28) suggested that root recovery experiments in soybean, pea, and snapbean allowed separation of H+ toxicity effects from Al3+ toxicity effects. Zhang et al. (178) showed that root regrowth after aluminum stress could be used to improve aluminum tolerance in triticale (Triticosecale spp.).

Genetics

Aluminum tolerance is a heritable trait in sorghum (179), barley (180), wheat (181,182), rice (Oryza sativa L.) (183), soybean (184), and Arabidopsis thaliana (185). With sorghum, Magalhaes (cited in 179) has found a pattern of inheritance of aluminum tolerance that is consistent with a single locus. With barley, Tang et al. (180) confirmed that aluminum tolerance segregation in F2 genotypes was due to a single gene, Alp, and they proposed the use of molecular markers in selection of aluminum tolerance in barley genotypes without the need for field trials, soil bioassays, or solution culture tests. In wheat, controversy exists over the number and location of genes that are involved in aluminum tolerance (181,182). In rice, nine different genomic regions on eight chromosomes have been associated with genetic control of plant response to aluminum, indicating that aluminum tolerance is a multigenic trait (183). Similarly, with soybean, aluminum tolerance is likely to be governed by 3 to 5 genes (184). In Arabidopsis, two quantitative trait loci occurring on two chromosomes could account for 43% of total variability in aluminum tolerance among a recombinant inbred population (185). A recent review of genetic analysis of aluminum tolerance in plants is found in Kochian et al. (179).