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B. F. CARWR AND J. D. OWNBY
erates what is already a natural process (Helyar and Porter, 1989). Certain wheatland soils in the Great Plains were acidic when first cultivated a century ago and
have acidified further with nitrogen fertilization and crop removal.
Two critical questions emerge in the development of an economically viable
wheat production system: To what degree should corrective additions of lime or
other basic materials be used to ameliorate the effects of acidification, and to what
degree should genetic alteration be used to overcome the limitations of reduced
fertility in acid soils? Answers to these questions are certainly not mutually exclusive, nor should either be expected to provide the sole solution. Scott and Fisher
(1989) argue with supporting evidence that tolerant cultivars are just as critical to
managing acid wheatland soils as lime application. Grain yields can be sustained
at lower pH and, therefore, with lower lime inputs by choosing tolerant cultivars.
Obviously, there are two parts (management and tolerance) to the equation for
achieving sustainable production in acid soils, and they must be properly balanced
as inputs change and genotypes improve.
Other factors enter into the equation because lime application is not the only
means for correcting acidification. Helyar and Porter ( 1989) prescribe more effective management of the nitrogen and carbon cycles as environmentally sound alternatives. For example, Fisher and Scott ( l993b) recommend earlier fall planting
of wheat in acid soils of southern NSW to minimize acidification caused by the
loss of added nitrate. Earlier planting also allows the wheat crop a “jump-start”
before waterlogging conditions occur in the winter. Researchers are also investigating other management alternatives which ameliorate the effects of acidification
in the highly sensitive seedling stages. Phosphorus precipitates in an unavailable
form in acid soils as it reacts with Al. The flip side to this reaction is that Al also
precipitates in the seedling root zone, which may explain why wheat yields can be
sustained when phosphate fertilizer is applied with the seed at planting (Boman
et al., 1991). Phosphorus provides more protection from Al if banded in the seedling root zone as opposed to broadcast application, even when the soil test index
indicates adequate P for wheat production. Boman el al. (1992) found rates of 34
and 67 kg ha - I of P20ssufficient to sustain normal grain and forage yields from
liming. Returns on investment make banded P applications highly economical for
producers who lease land on a yearly basis. Still the most economical approach to
sustained yields in acid soils over several years is to apply lime with adequate P
and to plant tolerant cultivars.
Tolerance to acid soils bears at least some concern in almost every agronomic
plant grown throughout the world. Wheat is no exception and has actually cata-
ACID SOIL TOLERANCE IN WHEAT
lyzed global concerns of the phytotoxic elements associated with acid soils.
Awareness of soil acidity has increased dramatically in wheat production areas
historically regarded as pH safe. Locations undergoing increasing acidification
include the wheat belts of the United States, Canada, Australia, and South Africa.
This increased awareness has led to advances in our knowledge of the biochemical
and genetic basis of adaptations to acidic wheatland soils. For example, a significant component of both Al and Mn tolerance now appears to be chelation of these
phytotoxic metals with organic acids. In the case of Al this probably occurs in the
apoplast or rhizosphere as part of an Al exclusion mechanism. There is also circumstantial evidence that constituents of the cell wall and plasmalemma are directly involved in exclusion; however, the details are not yet known. Technology
now exists to isolate and characterize proteins encoded by genes involved in A1
tolerance. In the next decade we expect that the nature of some of these proteins
and their functions will be revealed.
Although less is known about Mn tolerance mechanisms, the ability of the plant
to achieve a uniform distribution of leaf Mn, in sites apart from cell metabolism,
seems to be critical. Silicon and organic acids, including oxalate, are likely to play
important roles in tolerance. Manganese-tolerant and Mn-sensitive spring wheat
lines have been identified, and current assays should permit assessment of Mn
tolerance among winter wheats. The next step is to survey these cultivars to determine which biochemical and physiological parameters correlate with Mn tolerance in wheat.
Wheat researchers now have a better appreciation for the complexity of genetic
control of A1 and/or Mn tolerance. Recent work in locating Al tolerance genes to
chromosomes demonstrates that several genes can influence phenotypic expression and with different magnitude. These genes appear to be concentrated in the
A and D genomes. The specific genetic model used to describe inheritance of A1
tolerance in one series of progeny may not apply to another series, or even the
same series when challenged at a different level of stress. Gene expression may
be altered in different backgrounds, or possibly different genes are expressed at
different levels of stress. One common thread to genetic control of Al tolerance in
wheat is dominance expression in heterozygous materials; still, exceptions to that
trend are found, and the possible role epistasis plays in nonadditive gene action is
not well explored. While significant improvements have been made, particularly
for AI tolerance, by selection of major genes with easily identifiable effects, further advances will require manipulation of polygenes with much smaller effects,
perhaps too small to detect without the aid of markers. Further advances will also
be realized with due consideration to other yield-limiting factors inherent in an
acidic production system, such as improvement in phosphorus-use efficiency and
water-use efficiency of wheat.
Improved understanding of acid soil tolerance has not come without voids of
information. For example, linkage of physiological mechanisms of tolerance with
B. F. CARVER AND J. D.OWNBY
genetic mechanisms are lacking in wheat, but can be addressed with the recent
availability of near-isogenic stocks. The physiological characterization of tolerance genes from alien species is needed to verify uniqueness of control and to
warrant interspecific gene transfer. Gene location in other backgrounds exceeding
the tolerance level of Chinese Spring will provide evidence of allelic variation vs.
nonallelic variation for A1 tolerance. Genetic variation has not been described well
in wheat regarding allelic or nonallelic forms. Geneticists might focus on the B
genome to identify sources of tolerance to complement genes already present in
the A and D genomes; use of wheat-rye translocations offers one possible mechanism. Finally, greater priority has been given to the genetic improvement of A1
tolerance without systematically weighing the agronomic and economic benefits
of A1 vs. Mn tolerance, at least in U.S. wheatland soils. These information gaps
provide incentive to further advance an already active but fragmented research
area, one that beseeches cooperation of specialists in soil fertility, stress physiology, wheat breeding, and cellular and molecular biology.
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