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Mitigation
of Mineral Deficiency Stress |
By Dr.s
Surya Kant and Uzi Kafkafi The Hebrew University |
1.
Avoidance of Nutrient Stress
The efficiency of fertilizers might be lower than fifty per cent either due to mismanagement or due to reactions of the fertilizers with soil constituents. only a fraction of the fertilizer applied to the soil is taken up by the crop, the rest is either remains in the soil or lost through leaching, physical wash off, fixation by the soil, or release to the atmosphere through chemical and microbiological processes (Hera, 1996). Fertilizer losses especially of nitrogen could happen during the period of plant growth and lead to low yields than expected. This
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low uptake and
utilization of nutrients out of the added fertilizers results in lower
production and appearance of nutrient stress symptoms. The poor efficiency of
added fertilizers is mainly due to imbalance fertilization or improper
management practices. Critical information on the fertilization practices like:
method of fertilizer placement, time and rate of application and type of
fertilizers are essential for successful yields.
There are three
general main types of nitrogen fertilizers where the nitrogen in the fertilizer
is found in the chemical form of: ammonium, urea and nitrate. Nitrogen in
nitrate form fertilizer are soluble and highly mobile
in soil. They are prone to various kinds of losses: leaching beyond the root
zone in the light textured soils, denitrification in
water logged conditions or short time of over irrigation (Bar-Yosef and Kafkafi, 1972). Ammonium type fertilizers usually transform
in the soil within a week or two to nitrate form and the further consideration
is the same as for nitrate fertilizers.
Phosphate fertilizers, which are highly
reactive, are fixed in soil and become immobile. Potassic
fertilizers are less mobile, since they are adsorbed on the clay complex. The
entire quantity of phosphate and potassic
fertilizers are, therefore, applied in one dose at the time of sowing in
annual crops. Whereas, split application of nitrogenous
fertilizers increases nitrate reductase activity,
uptake of nitrogen (Abrol, 1990), use efficiency (Destain et al., 1997) and grain yield (Sabbe and Batchelor, 1990).
Application of
fertilizers in narrow bands beneath and by the side of the crop rows i.e. band
placement is preferable when the crop needs initial good start; when the soil
fertility is low; when fertilizer materials react with soil constituents
leading to fixation (phosphorus) and where volatilization losses are high.
Addition of
organic matter to the soil at regular periods, helps
to maintain the buffer capacity of the soil to supply the essential elements
after its decomposition. Organic matter in soils can be regulated through
different agronomic practices such as application of compost, farmyard manure, vermicompost, green manure etc’.
2.
Management of Nutrient Stress
When the soil is deficient in a particular nutrient element and the crop grown on such soil show visible deficiency symptoms, there is urgent need to correct it to improve the crop production. Methods of correcting nutrient deficiencies vary according to agro-climatic regions, the socioeconomic situations of the region, the magnitude of disorder, and nutrient element involved. A generalized description of these methods is presented in Table 2. Use of nutrient-efficient cultivars in combination with fertilizers or amendments may be the best solution for correcting nutrient disorders in field crops, but this will vary according to situation. Fertilizer recommendations are usually based on the results of field trials in which crop response to various rates of fertilizer application is determined. Such response curves provide relationships between yield and the amount of fertilizer required for a particular crop grown in a specific agro-climatic region.
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Table
2. Methods
of correcting nutrient deficiency (ased on Fageria et al., (1997b) |
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|
Nutrient element |
Corrective
Measures |
|
Nitrogen |
Addition of organic matter to soil;
application of N fertilizer, including legume in crop rotation; use of foliar
spray of 0.25-0.5% solution of urea |
|
Phosphorus |
Application of amendments to maintain
soil reaction to near neutral in acidic soils; application of phosphorus
fertilizer |
|
Potassium |
Application of potassium fertilizers,
incorporation of crop residues, manures |
|
Calcium |
Liming (addition of CaCO3) of
acid soils; addition of gypsum or other soluble calcium source where lime is
not required |
|
Magnesium |
Application of dolomite limestone;
foliar application of magnesium sulfate or magnesium nitrate solutions |
|
Sulfur |
Use of fertilizers containing sulfur
such as ammonium sulfate; single super phosphate; gypsum or elemental sulfur
(in acidic soils only). |
|
Zinc |
Addition of zinc sulfate to soil;
foliar spray of 0.1-0.5% solution of zinc sulfate |
|
Iron |
Foliar spray of 2% iron sulfate or
0.02-0.05% solution of iron chelate; use of
efficient cultivars, fertigation with chelated iron |
|
Copper |
Soil application of copper source of
fertilizer or foliar spray of 0.1-0.2% solution of copper sulfate |
|
Boron |
Soil application of boron source or
foliar spray of 0.1-0.25% solution of borax, care not to exceed 0.5 ppm B in solution in irrigation |
|
Molybdenum |
Liming of acid soils; soil application
of sodium ammonium molybdate; foliar spray of
0.07-0.1% solution of ammonium molybdate |
|
Manganese |
Foliar application of 0.1% solution of
manganese sulfate |
In a standing crop
if the nutrient deficiency is confirm, the foliar application of selected nutrients
by means of spray is a quick way to get rid of stress symptoms and avoiding
yield loss. Foliar fertilization of macro and micronutrients is best practice
whenever nutrient uptake through the roots is restricted due to adverse growing
conditions (El-Fouly and El-Sayed,
1997), when topsoil is dry particularly in semiarid regions (Grundon, 1980), under saline soils (El-Fouly
and Salama, 1999), and when root activity decreases
during reproductive stage (Ikeda et al., 1991). In calcareous soils iron
availability is generally very low and chlorosis is
quite common, foliar spraying under these conditions is very beneficial (Horesh and Levy, 1981). For example, iron chlorosis was corrected by foliar application of 0.1%
aqueous solution of Fe applied at 2 Kg Fe ha-1, either as iron
citrate or as iron sulfate + 400 g ha-1 citric acid (0.02%), in four
equal splits at 30, 45, 60 and 75 days after emergence in groundnut (Singh and
Joshi, 1997). Manganese deficiency in such soils is also widespread and two or
more sprays may be required within a growing season (Gettier
et al., 1985). Not all crops respond equally to foliar nutrition and the
nutrient elements differ in their rate of uptake through foliage and the degree
of mobility within the plant once absorbed (Table 3).
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Table
3.
General absorption and mobility rankings for foliar applied nutrients (from C
F A, 1985). |
|
|
Absorption |
Mobility |
|
Rapid Urea
Nitrogen, Potassium, Zinc Moderate Calcium, Sulfate, Manganese, Boron Slow Magnesium, Copper, Iron, Molybdenum |
Urea Nitrogen, Potassium, Phosphorus,
Sulfate Partially Zinc, Copper, Manganese, Boron,
Molybdenum Immobile Iron, Calcium, Magnesium |
|
Fig.2. Possible mechanisms in genotypic
differences for nutrient efficiency (Marschener,
1977b) |
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Nutrient
concentration and uptake by different plant genotypes are the most important
criteria for identifying the existing genetic specificity of plant nutrition (Saric, 1987). The tolerance in a given plant species or
genotype to nutrient stress is closely related to its nutrient use efficiency.
Genotypic differences in nutrient use efficiency are linked with root
nutrient acquisition capacity, or with utilization by
the plant, or both. Various plant species have developed morphological and/or
physiological mechanisms to improve acquisition and use efficiency of mineral
nutrients when grown on poor, infertile soils (Romeheld,
1998). By such mechanisms plants achieve nutrient deficiency stress tolerance.
However, sometimes the ability to adapt to stress is limited, depending on the
intensity and duration of stress. For example, some nutrient stress factors,
when operating for a long period, can result in inhibited growth and root
physiological processes responsible for nutrient uptake with the consequence of
an impaired capability for adaptation and stress tolerance.
For a given
genotype, nutrient use efficiency is reflected by the ability to produce a high
yield in a soil that is limiting in one or more mineral nutrients for a
standard genotype (Graham, 1984). The efficiency should be compared under
stress and adequate nutrient supply to verify plant species or genotypic
differences in nutrient utilization under sub-optimal and optimal conditions.
The differential response of genotypes to nutrient stress is related to uptake,
transport and utilization pattern of nutrients within plants (Fig.2). Isolation
of intra-species differences in capacity for growth under specific nutritional
conditions, particularly under nutritional stress is a critical aspect in
nutritional plant genetics. A wide range of morphological, anatomical and
physiological plant traits can be responsible for variations in response to
nutrient stress within a plant species. Such traits can function either within
or outside the plant to affect nutrient availability, absorption, translocation
or utilization. Some physiological and morphological plant features controlling
plant resistance to nutritional stress are given in Table 4.
The
acquisition of nutrients by plant roots plays the most important role in
tolerance to nutrient stress. Genotypes can differ widely in both the affinity
of uptake and the threshold concentration, for example, the uptake of
phosphorus in corn inbred lines (Schenk and Barber, 1979). The differences in
their ability to grow in soils of low phosphorus status might be attributed to
several factors including differential influx rates, root length/shoot weight
ratios (Fohse et al., 1988) or root system morphology and
root hair density (Randall, 1995). Among the more important factors is the role
of root exudates in making available soil P 'fixed' in forms such as iron or
aluminum phosphate. White lupin and pigeon pea are
well adapted to acidic P-deficient soils (Hocking et al., 1997). White lupin forms develop proteoid
roots (bottlebrush-like clusters of rootlets covered with a dense mat of root
hairs) which are considered to be a response to low P availability (Marschner, 1986). Proteoid roots
of white lupin secrete citric acid which solubilizes 'fixed' P (Gardner et al., 1983; Gerke et al.,1994), thus
increasing P uptake by the plant. For pigeon pea, secretion of piscidic, malonic and oxalic
acids appears to be the mechanism by which this species is able to release P
from iron and aluminum phosphates (Otani et al.,
1996). Screening of tolerant Medicago
sativa and Lablab purpureus germplasm
under phosphorus stress condition was carried out by Mugwira
and Haque (1993a and b) based on their root and shoot
growth.
Genetic
differences in K uptake and utilization efficiency in field crop cultivars have
also been observed (Glass et al., 1981). Growth response of 20 wheat genotypes
were compared by Gill et al. (1997) under deficient (0.6 mM)
and adequate (3.0 mM) K levels in solution cultures.
Substantial and significant differences due to K-stress were obvious among
genotypes for shoot dry weight, relative reduction in shoot dry weight, root
dry weight, and root/shoot ratio. Rengel (1997)
screened wheat genotypes tolerant to Zn and Mn stress. Wheat genotypes tolerant
to Zn deficiency released greater amounts of phytosiderophore,
2-deoxymugineic acid, than the sensitive genotypes. In addition, Zn deficiency
increased the numbers of fluorescent pseudomonas in rhizosphere
of all wheat genotypes tested, but the effect was particularly obvious for
genotypes tolerant to Zn deficiency. Under Mn stress, wheat genotypes tolerant
of Mn deficiency had a greater ratio of Mn-reducers to Mn-oxidizers in the rhizosphere compared to sensitive genotypes. Mn-efficient
durum wheat genotypes had greater Mn uptake from Mn deficient soil, produced
higher grain yield, relative grain yield and above ground biomass and generally
maintained higher seed Mn concentration (Khabaz-Saberi
et al., 1997).
The Fe efficient
groundnut cultivar 'ICGV-86031' released more hydrogen ions and reductants from its roots under iron deficiency stress
treatment, maintained higher leaf Fe and had less chlorosis
than the Fe-inefficient cultivar 'TCGS-37' (Reddy et al., 1997). There
are also large differences in tolerance to Fe deficiency among field grown
wheat genotypes (Hanson et al., 1996).
Genetic control of
Cu efficiency has been located on the long arm of the rye chromosome 5 and has
been applied to wheat breeding for CU efficiency (Podlesak
et al., 1990).
Similar, increase
in Cu efficiency was observed by Graham (1984) in spring form of rye (Secale cereale L.)
and genetically manipulated wheat. Wheat genotypes differ in tolerance to Zn
deficiency (Graham et al., 1992; Cakmak et
al., 1996a), with durum wheat being generally less tolerant than bread
wheat (Rengel and Graham, 1995; Cakmak
et al., 1996). Tolerance of bread wheat genotypes to Zn and Fe
deficiency was expressed in shoot dry weight and relative shoot growth as
compared with to sensitive genotypes (Rengel and Romheld, 2000).
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Cereal species, as
well as genotypes of a given cereal greatly differ in adaptation to
Zn-deficient conditions. The susceptibility of cereals to Zn deficiency was
found to decline in the order: durum wheat > oat > bread wheat >
barley > triticale > rye. Among the cereals, rye
showed an exceptionally high Zn efficiency. Biomass production and grain yield
of rye was not affected under severe Zn-deficient conditions (Cakmak et al., 1997b). Several mechanisms constitute
the physiological bases of Zn efficiency. These are enhancement of root growth
(Dong et al., 1995), root uptake and root-to-shoot translocation of Zn (Cakmak et al., 1996a; Rengel
and Graham, 1996), release of Zn-mobilizing phytosiderophores
from roots (Cakmak et al., 1996b; Erenoglu et al., 1996) and internal utilization of
Zn (Cakmak et al., 1996b; Rengel,
1995). Use of disomic wheat-rye addition lines (Triticum aestivum
L., cv. Holdfast-Secale
cereale L., cv.
King-II) increases Zn efficiency (Table 5). The addition of rye chromosomes,
particularly 1R, 2R and 7R, into Holdfast reduced the severity of deficiency
symptoms (Cakmak et al., 1997a). Transgenic
plants have improved nutrient efficiency and resistant to nutrient stress. This
was found for copper in wheat (Graham et al., 1987); boron and phosphorus in
tobacco (Brown et al., 1999; Lopez-Bucio et al.,
2000).
The molecular
approach to breeding of mineral deficiency resistance and mineral efficiency is
now receiving increasing attention. QTLs (quantitative trait loci) controlling
Phosphorus efficiency were identified in rice and maize. Transgenic tobaccos overexpressing ferritin in the
plastids or in the cytoplasm resulted in higher leaf concentration of iron,
manganese and zinc (see #4791 in this site reference database). Tobacco
transformation over expressing the antioxidant superoxide
dismutase had greater manganese deficiency resistance
(#4512).
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