Bạn đang xem bản rút gọn của tài liệu. Xem và tải ngay bản đầy đủ của tài liệu tại đây (3.5 MB, 343 trang )
ADVANCES IN CHLORIDE NUTRITION
underground water table, the draining water may create a continuous wet zone with
the groundwater level In that case, the upward ﬂow of capillary water leads to
salinization of the soil surface as a result of water evaporation with chloride precipitation on the soil surface.
In drip irrigation, the salt distribution pattern depends on the rate of evaporation
from the soil surface (Yaron et al., 1973), the water uptake by the plant, the location of the wetting front, the total amount of applied irrigation water, and the distance between drip lines. As the amount of applied water increases, more salts are
leached below the drip line and there is a larger salt accumulation on the soil surface between the irrigation lines.
Chloride concentration in the soil solution increases as more water is taken up
by the plant and the soil moisture level approaches the permanent wilting point.
Plants can tolerate water with a high salt concentration when the soil moisture level is high (Rhoades, 1993) and when the high salt concentration zone is located in
deeper soil layers with lower root density. Therefore, classiﬁcations of water quality should also consider the effects of irrigation practices and plant root distribution.
B. MONITORING OF CHLORIDE CONCENTRATION IN SOIL
The correct management of irrigation requires periodic monitoring of the concentration of the soil solution in the root zone. The salt concentration must remain
below a given threshold value, according to the sensitivity of the crop to salinity
and particularly to chloride (see Table V).
The methods used for monitoring soil salinity were reviewed by Rhoades and
Oster (1986). The total concentration of soluble salts in the soil solution is generally estimated by determining the EC of the saturated paste extract or by sampling
the soil solution. In most cases, chloride is the main anion present in the soil solution. The chloride concentration can be determined directly in the saturated soil extract or in the soil solution extracted with suction cups or with gypsum block sensors. Chloride testing methods have been reviewed by Johnson and Fixen (1990).
The reliability of the results is inﬂuenced by the soil variability, the position of
the soil solution suction cups relative to the pattern of water and salt accumulation
in the soil proﬁle, and sensors type (Rhoades and Miyamoto, 1990). The number
of samples and their spatial distribution will affect the reliability of the ﬁeld salinity level determination.
C. FERTILIZATION UNDER CONDITIONS
OF CHLORIDE SALINITY
With the increasing use of saline and recycled sewage water for agriculture, fertilizer application under saline conditions has become a subject of considerable in-
GUOHUA XU ET AL.
terest (Feigin, 1985). Sodium chloride salinity disrupts mineral nutrition acquisition by plants in two ways (Grattan and Grieve, 1992): (1) total ionic strength of
the soil solution, regardless of its composition, can reduce nutrient uptake and
translocation; and (2) uptake competition with Naϩ and Cl ions can reduce nutrient availability. These interactions may lead to Naϩ-induced Ca2ϩ and/or Kϩ
deﬁciencies (Volkmar et al., 1998) and Cl-induced inhibition of nitrate uptake
(Kafkaﬁ et al., 1982). It was postulated that the salinity tolerance of crops can be
improved by the suitable use of nutrients (Kafkaﬁ, 1987).
Most of the reported studies on NaCl salinity effects do not separate the effects of
Naϩ from those of Cl (Volkmar et al., 1998). A low concentration of Cl salts is beneﬁcial (Fixen et al., 1986). In the wet volume of irrigated soils used to grow regular
ﬁeld crops such as tomatoes and melons, EC values of about 3.0 dS/m in the soil solution, with chloride salts as the dominant component, are common (Kafkaﬁ et al.,
1992). Chloride concentrations above about 10 mmol/liter in the irrigation water
are generally considered problematic for plant growth (Ayers and Westcott, 1985).
Salinization with NaCl or KCl inhibits the net uptake of nitrate in citrus (Cerezo et
al., 1997) and causes nitrogen deﬁciency (Embleton et al., 1978). Nitrate competes
with chloride for uptake by the plant, as discussed in Section IIIE; therefore, when
the irrigation water contains nitrate at about 8–16 mmol/liter, even sensitive plants
like avocado can survive at chloride concentrations of 8–16 mmol/liter (Fig. 3).
High potassium fertilization might enhance the capacity for osmotic adjustment
of plants growing in saline habitats (Cerda et al., 1995), as potassium is the most
abundant cation in the cytoplasm of glycophytes (Marschner, 1986). In spinach,
higher Kϩ requirements are needed for shoot growth under high salinity than under low salinity conditions (Chow et al., 1990). Differences in salt tolerance among
maize varieties appear to be related to higher Kϩ ﬂuxes and cytoplasmic concentrations on the one hand and lower Naϩ and Cl ﬂuxes an cytoplasmic concentrations on the other (Hajibagheri et al., 1989). Potassium uptake is greater in the high
salt-tolerant group of barley cultivars than in salt-sensitive ones (Sopandie et al.,
1993). However, an external Kϩ supply is not required for root growth of castor
bean (Ricinus communis L.) under saline conditions ( Jeschke and Wolf, 1988). Increasing the Kϩ supply in the rooting media of maize did not alleviate the growth
reduction imposed by treatment with NaCl at 50 mmol/liter (Cerda et al., 1995).
Tip-burn symptoms in Chinese cabbage, induced by salinity with NaCl and CaCl2,
was not alleviated by the addition of KNO3 (Feigin et al., 1991).
The addition of adequate P can also be helpful in alleviating salt stress (Champagnol, 1979; Awad et al., 1990). A positive effect on P on the yield of foxtail millet and clover grown in a saline soil was reported (Ravikovitch and Yoles, 1971).
As crops can differ greatly in their response to nutrition management under different combinations of environmental salinity (Feigin et al., 1991), speciﬁc information on the behavior of crops under different situations of Cl salinity is needed for
the optimal fertilization management of speciﬁc crops.
ADVANCES IN CHLORIDE NUTRITION
D. IRRIGATION AND LEACHING OF CHLORIDE SALTS
IN THE ROOT ZONE
The use of water containing chloride must be accompanied by appropriate practices to keep Cl levels in the soil within the limits of crop tolerance. The amount
of supplementary Cl salt added to the soil depends on the salt concentration in the
water, the evaporation level and the amount of irrigation water (which depends on
the physiological development of the plant). With 500 mm of irrigation water containing Cl at 100 –200 mg/liter (low to medium level of salinity, see Table I), the
applied Cl reaches 500 –1000 kg/ha. This amount of Cl is equivalent to KCl fertilization of 1000 –2000 kg/ha. In ﬁeld practice, the recommended range of KCl
fertilization is 75–150 kg/ha for ﬁeld crops and 300–500 kg/ha for horticultural
crops. This suggests that the addition of Cl in KCl fertilizer is relatively safe for
most agricultural crops, especially when the rainfall during the rainy season is capable of leaching the excess Cl accumulated during fertilization and irrigation.
Irrigation with saline water is managed by an excess of irrigation to meet the
leaching requirement for avoiding salt accumulation in the crop root zone
(Richards, 1954). When plants are present, however, there is a risk that the advantages gained from salt leaching may be lost with the onset of temporary oxygen shortage due to water-logging (Stevens and Harvey, 1995). The ability of
grapevine roots to exclude sodium and chloride from the leaf was strongly reduced
by a short period of waterlogging (West and Taylor, 1984; Stevens and Harvey,
1995). In a river land of southern Australia, the EC of irrigation water during the
period 1985–1990 was less than 0.5 dS/m; however, grapevines suffered salinity
damage because of excessive irrigation. The excess water drained to an aquitard
just below the root zone and formed a temporary water table that mobilized the
previously leached salts back into the root zone as a result of the capillary rise of
the water (Stevens and Harvey, 1995).
The irrigation system inﬂuences the distribution of salt in the soil’s proﬁle and
surface. Keller and Bliesner (1990) presented a detailed calculation of the efﬁciency of chloride leaching by different irrigation methods. In drip irrigation, the
water is applied at short intervals so that the application of minimum leaching doses and the relatively small change in the soil’s water content keep the salinity of
the soil close to that of the irrigation water. In drip irrigation the leaching dose
(LRt) required to wash salts out of the root zone is deﬁned as the ratio between the
water height applied for leaching and the irrigation water height applied for satisfying crop and leaching demands. The equation is simply expressed as: LRt ϭ
ECw /2ECemx, where ECw is the EC of the irrigation water and ECemx is the maximum EC value of the saturated soil extract at which the crop can survive.
In sprinkler or surface irrigation the water is applied at longer intervals, during
which the salt concentration of the soil solution gradually increases. Before the
next irrigation, salt concentrations on the soil surface reach relatively high values.
GUOHUA XU ET AL.
The equation for calculating the leaching requirement dose is: LRt ϭ ECw /(5ECe
Ϫ ECw), where ECe is the mean EC of the saturated soil extract at which no yield
reduction occurs (Keller and Bliesner, 1990).
E. SPRINKLER IRRIGATION AND CHLORIDE-INDUCED
Chloride can be absorbed directly by crop leaves and can cause foliar injury
when its concentration in the sprinkler water is high (Maas et al., 1982). Leaf
scorching due to excessive Cl accumulation in the leaves varies among different
species and depends on leaf properties and on the rate of Cl absorption by the
leaves. Temperature, relative humidity, and water stress all have marked effects on
the leaf injury. Absorption of Cl continues as long as the leaf is wet. Evaporation
from the leaf surface increases the salt concentration on the leaf and consequently also the leaf scorching level.
Deciduous trees, such as almond, apricot, and plum, absorb Naϩ and Cl readily through the leaves, and partial leaf abscission occurs after a 50-hr sprinkler irrigation with water containing CaCl2 or NaCl at a concentration of 10 mmol/liter
(Ehlig and Bernstein, 1959). The crop leaf Cl concentrations causing leaf injury in
plum, almond, and orange are 4.3–7.1, 6.4–10.6, and 7.1–10.6 mg/g DM, respectively. No visual foliar injury was observed in grapes sprinkled with water
containing Cl at 5 or 10 mmol/liter (Francois and Clark, 1979). In avocado, where
the thick waxy layer of the leaves limits the absorption of ions present in the sprinkling water, chloride accumulation in the leaves is very low and no visual injuries
are observed (Ehlig and Bernstein, 1959). Therefore, although avocado is known
to be sensitive to salt concentration in the growing medium, there is no risk of direct foliar absorption of chloride because of the leaf surface characteristics. Chloride accumulation in crop leaves depends mainly on the time of watering (Francois and Clark, 1979). Therefore, rootstocks known to limit chloride absorption by
the roots are not suitable when sprinkler irrigation is used and they do not avoid
chloride accumulation in the leaves.
Field and vegetable crops are not especially sensitive to salt accumulation in the
leaves (Ehlig and Bernstein, 1959). Strawberry is highly sensitive to chloride in
the soil solution, but is less affected by salt absorption through leaves (Ehlig,
1961). The rate of foliar absorption of chloride increases in the following order:
sorghum ϽϽcotton, sunﬂowerϽcauliﬂowerϽsesame, alfalfa, sugar beetϽbarley,
tomatoϽpotato, safﬂower (Maas et al., 1982). However, this order does not apply
to foliar injury. The relative values of crop sensitivity to foliar injury due to chloride in the sprinkling water are summarized in Table VIII. Because both crop and
environmental conditions inﬂuence the injury level, these data constitute only a
guideline to irrigation during daytime hours.
ADVANCES IN CHLORIDE NUTRITION
Crop Sensitivity to Foliar Injury due to Chloride
in Sprinkler Irrigation Watera
Crops exhibiting foliar injury
Almond, apricot, citrus, plum
Grape, pepper, potato, tomato
Alfalfa, barley, corn, cucumber, safﬂower,
Cauliﬂower, cotton, sugar beet, sunﬂower
on Maas et al. (1982).
Sprinkler irrigation of crops that are less sensitive to chloride is possible provided that steps are taken to avoid or minimize foliar injuries. Such measures might
include the use of mobile sprinklers, uniform water distribution, night irrigation,
and the scheduling of longer intervals between irrigations (Maas, 1985).
Chloride anions are hardly sorbed to soil particles and are easily leached in soil
proﬁles. In acid soils containing variable-charge clays, a slight speciﬁc sorption of
chloride is observed.
The crop response to Cl varies among genera, species, and cultivars. The lowest critical Cl concentration for plants below which response to Cl addition is observed ranges between 0.1 and 6 mg/g DM or between about 0.03 and 17 mmol/
liter of chloride on the basis of the plant tissue water content. The normally nontoxic Cl concentrations in plants range from 1 to 20 mg/g. The Cl concentration
in the plant depends in part on the concentration of Cl in the external solution and
its ratio to other anions, particularly nitrate. Chloride compartmentation appears
to be highly regulated. In the chloroplast, the Cl concentration remains relatively
constant regardless of whether Cl in the soil solution is deﬁcient or excessive.
Chloride is required for photosynthesis, charge compensation, and osmoregulation of the whole plant, as well on a single cell basis as in stomatal guard cells.
Palms and coconuts use Cl for charge balance in the guard cells. Relatively large
amounts of Cl are essential for some crops, such as kiwifruit and sugar beet.
Diagnosis of salt toxicity in plants must distinguish between the effects of chloride and those of the accompanying sodium cation. The overall tolerance to high
GUOHUA XU ET AL.
concentrations of external chloride is due to the ability of the plant to limit Cl uptake by the roots and its transport to the shoots. Accumulation of Cl in the leaves
depends both on its rate of uptake and translocation from the roots to the leaves.
In most crops, the accumulation of chloride in the leaves is controlled by the rootstock. Chloride-sensitive cultivars accumulate an excessive amount of Cl in the
shoots and tolerant cultivars restrict Cl transport to the shoots by a mechanism that
resides in the root. The level of accumulated Cl in the plant should not be regarded as the sole criterion of crop tolerance to chloride.
Ammonium stimulates Cl accumulation in plants. Nitrate can prevent Cl toxicity
at a concentration of up to 16 mmol Cl/liter in the soil solution. A model for the regulation of Cl inﬂux suggests that both negative feedback effects from vacuolar
3 /Cl) or total anion concentrations and external NO3 inhibition of Cl inﬂux at
the plasmalemma may be operating. These combined effects serve to discriminate
against Cl accumulation, favoring NOϪ
3 uptake and its subsequent metabolism. The
uptake interaction between chloride and phosphorus appears to be complex. Phosphate uptake is stimulated when chloride concentrations in the external solution are
low and suppressed when they are high. High levels of NaCl reduce Ca2ϩ and Kϩ
in the roots and leaves. The interaction of chloride with other plant nutrients needs
The mechanisms of the effects of Cl on foliar disease infection are not well understood. The possibility that both climatic and biological factors may interact with
the plant response to Cl makes it difﬁcult to interpret plant and soil diagnostic data,
except in cases of extreme Cl deﬁciency in the soil.
Irrigation water containing Cl at less than 150 mg/liter, with ECs in the range
of 1–3 dS/m, can be used for most crops, provided that management practices are
taken into consideration. The main fertilizers containing chloride are potassium
chloride and ammonium chloride. When the annual application rates of these fertilizers supply less than 140 kg Cl/ha, no negative effects on crop growth or yield
Adler, P. R., and Wilcox, G. E. (1995). Ammonium increases the net rate of sodium inﬂux and partitioning to the leaf of muskmelon. J. Plant Nutr. 18, 1951–1962.
Al-Harbi, A. R. (1995). Growth and nutrient composition of tomato and cucumber seedlings as affected by sodium chloride salinity and supplemental calcium. J. Plant Nutr. 18(7), 1403 –1416.
Ali, I. A., Kafkaﬁ, U., Yamaguchi, S., and Inanaga, S. (1998). Response of oilseed rape plant to low
root temperature and nitrate:ammonium ratios. J. Plant Nutr. 21(7), 1463 –1481.
Anderson, C. A., and Steveninck, R. F. M. (1987). Accumulation and subcellular distribution of sodium and chloride and potassium ions in lucerne populations differing in salt tolerance. Aust. Salinity Newsl. 15, 74 –75.
Andersson, B., Critchley, C., Ryrie, I. J., Jansson, C., Larsson, C., and Anderson, J. M. (1984). Modiﬁcation of the chloride requirement for photosynthetic O2 evolution. FEBS Lett. 168, 113 –117.
ADVANCES IN CHLORIDE NUTRITION
Arnon, D. I., and Whatley, F. R. (1949). Is chloride a coenzyme of photosynthesis? Science 110, 554–556.
Ashraf, M., and Fatima, H. (1995). Responses of salt tolerant and salt sensitive lines of safﬂower
(Carthamus tinctorius L.) to salt stress. Acta Physiol. Plant. 17(1), 61–70.
Ashraf, M., and O’Leary, J. W. (1949). Ion distribution in leaves of varying age in salt-tolerant lines of
alfalfa under salt stress. J. Plant Nutr. 17(8), 1463 –1476.
Awad, A. S., Edwards, D. G., and Campbell, L. C. (1990). Phosphorus enhancement of salt tolerance
of tomato crop. Crop Sci. 30, 123 –128.
Ayers, R. S., and Westcott, D. W. (1985). Water quality for agriculture. Irrig. Drain. Pap. 29, Rev. 1.
Baianu, I. C., Critchley, C., Govindjee, and Gutowsky, H. S. (1984). NMR study of chloride ion interactions with thylakoid membranes. Proc. Natl. Acad. Sci. U.S.A 81, 3713 – 3717.
Banuls, J. E., and Primo-Millo, E. (1992). Effects of chloride and sodium on gas exchange parameters
and water relations of citrus plants. Physiol. Plant. 86, 115 –123.
Banuls, J. E., and Primo-Millo, E. (1995). Effects of salinity on some citrus scion rootstock combinations. Ann. Bot. (London) [N.S.] 76(1), 97–102.
Banuls, J. E., Legaz, F., and Primo-Millo, E. (1990). Effect of salinity on uptake and distribution of
chloride and sodium in some citrus scion-rootstock combinations. J. Hortic. Sci. 65, 715 –724.
Banuls, J. E., Legaz, F., and Primo-Millo, E. (1991). Salinity-calcium interactions on growth and ionic concentration of citrus plants. Plant Soil 133(1), 39 – 46.
Banuls, J. E., Serna, M. D., Legaz, F., Talon, M., and Primo-Millo, E. (1997). Growth and gas exchange
parameters of citrus plants stressed with different salts. J. Plant Physiol. 150(1–2), 194 –199.
Bar, Y., and Kafkaﬁ, U. (1992). Nitrate induced iron-deﬁciency chlorosis in avocado rootstocks and its
prevention by chloride. J. Plant Nutr. 15(10), 1739 –1746.
Bar, Y., Apelbaum, A., Kafkaﬁ, U., and Goren, R. (1996). Polyamines in chloride stressed citrus plants:
Alleviation of stress by nitrate supplementation via irrigation water. J. Am. Soc. Hortic. Sci.
Bar, Y., Apelbaum, A., Kafkaﬁ, U., and Goren, R. (1997). Relationship between chloride and nitrate
and its effect on growth and mineral composition of avocado and citrus plants. J. Plant Nutr. 20,
Bell, P. F., Vaughn, J. A., and Bourgeois, W. J. (1997). Leaf analysis ﬁnds high levels of chloride and
low levels of zinc and manganese in Louisiana citrus. J. Plant Nutr. 20(6), 733 –743.
Beringer, H., Hoch, K., and Lindauer, M. G. (1990). Source:sink relationship in potato as inﬂuenced
by potassium chloride or potassium sulfate nutrition. In “Plant Nutrition: Physiology and Application” (M. L. van Beusichem, ed.), pp. 639 – 642. Kluwer Academic Publishers, Dordrecht, The
Bernstein, L., Ayers, A. D., and Wadleigh, C. H. (1951). The salt tolerance of white potatoes. Proc. Am.
Soc. Hortic. Sci. 57, 231–236.
Bernstein, L., Fireman, M., and Reeve, R. C. (1955). Control of salinity in the Imperial Valley, California. U.S. Agric. Res. Serv., ARS ARS-41-4.
Bernstein, L., Ehlig, C. F., and Clark, R. A. (1969). Effect of grape rootstocks on chloride accumulation in leaves. J. Am. Soc. Hortic. Sci. 94, 584 – 590.
Bingham, F. T., Fenn, L. B., and Oertli, J. J. (1968). A sand culture study of Cl toxicity to mature avocado trees. Soil Sci. Soc. Am. Proc. 32, 249 –252.
Bishop, A. A., Walker, W. R., Allen, N. L., and Poole, G. J. (1981). Furrow advance rates under surge
ﬂow systems. J. Irrig. Drainage Div., Am. Soc. Agric. Eng. 107 (IR3), 257–264.
Bonneaux, X., Boutin, D., Bourgoing, R., and Sugarianto, J. (1997). Sodium chloride, an ideal fertilizer for coconut palms in Indonesia. Plant. Rech. Dev. 4(5), 336 – 346.
Borggaard, O. K. (1984). Inﬂuences of iron oxides on nonspeciﬁc anion (chloride) adsorption by soils.
J. Soil Sci. 35, 71–78.
Boursier, P., Lynch, J., Lauchli, A., and Epstein, E. (1987). Chloride partitioning in leaves of salt
stressed sorghum, maize, wheat and barley. Aust. J. Plant Physiol. 14(4), 463 – 473.
GUOHUA XU ET AL.
Braconnier, S., and d’Auzac, J. (1990). Chloride and stomatal conductance in coconut. Plant Physiol.
Biochem. 28, 105 –112.
Broyer, T. C., Carlton, A. B., Johnson, C. M., and Stout, P. R. (1954). Chloride–a micronutrient element for higher plants. Plant Physiol. 29, 526 – 532.
Buwalda, J. G., and Smith, G. S. (1991). Inﬂuence of anions on the potassium status and productivity
of kiwifruit (Actinidia deliciosa) vines. Plant Soil 133, 209 –218.
Callan, N. W., and Westcott, M. P. (1996). Drip irrigation for application of potassium to tart cherry. J.
Plant Nutr. 19(1), 163 –172.
Cao, W., and Tibbitts, T. W. (1993). Study of various ammonium/nitrate mixtures for enhancing growth
of potatoes. J. Plant Nutr. 16(9), 1691–1704.
Cerda, A. J., Pasrdines, M., Botella, A., and Martinez, V. (1995). Effect of potassium on growth, water relations and the inorganic and organic solute contents for two maize cultivars grown under
saline conditions. J. Plant Nutr. 18(4), 839 – 851.
Cerezo, M., Garcia-Agustin, P., Serna, M. D., and Primo-Millo, E. (1997). Kinetics of nitrate uptake
by citrus seedlings and inhibitory effects of salinity. Plant Sci. 126, 105 –112.
Champagnol, F. (1979). Relationship between phosphate nutrition of plants and salt toxicity. Phosphorus Agric. 76, 35 – 44.
Chapman, H. D., and Liebig, G. F. (1940). Nitrogen concentration and ion balance in relation to citrus
nutrition. Hilgardia 13, 141–173.
Chartzoulakis, K. S. (1991). Effects of saline irrigation water on germination, growth and yield of
greenhouse cucumber. Acta Hortic. 287, 327– 334.
Chien, C. T., Shetty, K., Mortimer, M., and Orser, C. S. (1991). Calcium induced salt tolerance in Rhizobium leguminosarum biovar viciae strain C1204b. Microbiol. Lett. 83(2), 219 –224.
Chow, W. S., Ball, M. C., and Anderson, J. M. (1990). Growth and photosynthetic responses of spinach
to salinity: Implications of Kϩ nutrition for salt tolerance. Aust. J. Plant Physiol. 17(5), 563 – 578.
Christensen, N. W., Taylor, R. G., Jackson, T. L., and Mitchell, B. L. (1981). Chloride effects on water potentials and yield of winter wheat infected with take-all root rot. Agron. J. 73, 1053 –1058.
Christensen, N. W., Roseberg, R. J., Brett, M., and Jackson, T. L. (1986). Chloride inhibition of nitriﬁcation as related to take-all disease of wheat. In “Special Bulletin on Chloride and Crop Production” (T. L. Jackson, ed.), No. 2, pp. 22– 39. Potash & Phosphate Institute, Atlanta, GA.
Churchill, K. A., and Sze, H. (1984). Anion-sensitive, Hϩ-pumping ATPase of oat roots. Plant Physiol. 76, 490–497.
Cline, R. A., and Bradt, O. A. (1980). The effect of source and rate of potassium on performance of
‘Concord’ grape vines grown on clay loam soils. J. Am. Soc. Hortic. Sci. 105(5), 650 – 653.
Cole, P. J. (1985). Chloride toxicity in citrus. Irrig. Sci. 6(1), 63 –71.
Coleman, W. J., Govindjee, and Gutowsky, H. S. (1987). The location of the chloride binding sites in
the oxygen-evolving complex of spinach photosystem II. Biochim. Biophys. Acta 894, 453 – 459.
Collins, J. C., and Abbas, M. A. (1985). Ion and water transport in seedlings of mustard (Sinapsis alba
L.). New Phytol. 99(2), 195 –202.
Corbett, E. G., and Gausman, H. W. (1960). The interaction of chloride and sulfate in the nutrition of
potato plants. Agron. J. 52, 94 – 96.
Cram, W. J. (1973). Internal factors regulating nitrate and chloride inﬂux in plant cells. J. Exp. Bot. 24,
Cram, W. J. (1983). Chloride accumulation as a homeostatic system: Set points and perturbations. The
physiological signiﬁcance of inﬂux isotherms, temperature effects and the inﬂuences of plant
growth substances. J. Exp. Bot. 34, 181–1502.
Cram, W. J. (1988). Transport of nutrient ions across cell membranes in vivo. Adv. Plant Nutr. 3, 1–54.
Critchley, C. (1985). The role of chloride in photosystem II. Biochim. Biophys. Acta 811, 33 – 46.
Davies, J. N., and Hobson, G. E. (1981). The constituents of tomato fruit—the inﬂuence of environment, nutrition, and genotype. Crit. Rev. Food Sci. Nutr. Chem. 15(3), 205 –280.
ADVANCES IN CHLORIDE NUTRITION
Deane-Drummond, C. E. (1986). A comparison of regulatory effects of chloride on nitrate uptake, and
of nitrate on chloride uptake into Pisum sativum seedlings. Physiol. Plant. 66(1): 115 –121.
Downton, W. J. S. (1985). Growth and mineral composition of the Sultana grapevine as inﬂuenced by
salinity and rootstock. Aust. J. Agric. Res. 36(3), 425 – 434.
Du, Z., Aghoram, K., and Outlaw, W. H., Jr. (1997). In vivo phosphorylation of phosphoenolpyruvate
carboxylase in guard cells of Vicia faba L. is enhanced by fusicoccin and suppressed by abscisic
acid. Arch. Biochem. Biophys. 337, 345 – 350.
Eaton, F. M. (1966). Chapter 8: Chlorine. In “Diagnostic Criteria for Plants and Soils” (H. D. Chapman, ed.), pp. 98–135. University of California, Riverside.
Ehlig, C. F. (1961). Salt tolerance of strawberries under sprinkler irrigation. Proc. Am. Soc. Hortic. Sci.
Ehlig, C. F., and Bernstein, L. (1959). Foliar absorption of sodium and chloride as a factor in sprinkler
irrigation. Proc. Am. Soc. Hortic. Sci. 74, 661– 670.
Embleton, T. W., Jones, W. W., and Platt, R. G. (1978). Leaf analysis as a guide to citrus fertilization.
Bull.—Univ. Calif. Div. Agric. Sci. 1879, 4 – 54.
Engel, R. E., Eckhoff, J., and Berg, R. (1994). Grain yield, kernel weight, and disease responses of winter wheat cultivars to chloride fertilization. Agron. J. 86, 891– 896.
Engel, R. E., Bruckner, P. L., Mathre, D. E., and Brumﬁeld, S. K. Z. (1997). A chloride deﬁcient leaf
spot syndrome of wheat. Soil Sci. Soc. Am. J. 61, 176 –184.
Engel, R. E., Bruckner, P. L., and Eckhoff, J. (1998). Critical tissue concentration and chloride requirements for wheat. Soil Sci. Soc. Am. J. 62, 401– 405.
Engvild, K. C. (1986). Chlorine-containing natural compounds in higher plants. Photochemistry 25,
Eshel, A., and Waisel, Y. (1979). Distribution of sodium and chloride in leaves of Suaeda monoica halophyte. Physiol. Plant. 46(2), 151–154.
Faiz, S. M. A., Ullah, S. M., Hussain, A. K. M. A., Kamal, A. T. M. M., and Sattar, S. (1994). Yield,
mineral contents and quality of tomato (Lycopersicon esculentum) under salt stress in a saline soil.
Curr. Agric. 18(1–2), 9 –12.
Feigin, A. (1985). Fertilization management of crops irrigated with saline water. Plant Soil 89, 285–299.
Feigin, A., Rylski, I., Meiri, A., and Shalhevet, J. (1987). Response of melon and tomato plants to chloride-nitrate ratio in saline nutrient solutions. J. Plant Nutr. 10(9 –16), 1787–1794.
Feigin, A., Pressman, E., Imas, P., and Miltau, O. (1991). Combined effects of KNO3 and salinity on
yield and chemical composition of lettuce and Chinese cabbage. Irrig. Sci. 12, 223 –230.
Felle, H. H. (1994). The Hϩ /ClϪ symporter in root-hair cells of Sinapsis alba. An electrophysiological study using ion-selective microelectrodes. Plant Physiol. 106(3), 1131–1136.
Fixen, P. E. (1987). Chloride fertilization. Crops Soils Manag. 39(6), 14 –16.
Fixen, P. E. (1993). Crop responses to chloride. Adv. Agron. 50, 107–150.
Fixen, P. E., Farber, B. G., Gelderman, R. H., and Gerwing, J. R. (1986). Role of Cl in maximum yield
environments: I. Evidence of yield response and Cl requirements. In “Special Bulletin on Chloride and Crop Production” (T. L. Jackson, ed.), No. 2, pp. 41– 51. Potash & Phosphate Institute,
Fixen, P. E., Gelderman, R. H., Gerwing, J. R., and Farber, B. G. (1987). Calibration and implementation of a soil Cl test. J. Fertil. Issues 4, 91– 97.
Flowers, T. J. (1988). Chloride as a nutrient and as an osmoticum. Adv. Plant Nutr. 3, 55 78.
Franỗois, L. E., and Clark, R. A. (1979). Accumulation of sodium and chloride in leaves of sprinklerirrigated grape cultivars. J. Am. Soc. Hortic. Sci. 104(1), 11–13.
Freney, J. R., Delwiche, C. C., and Johnson, C. M. (1959). The effect of chloride on the free amino
acids of cabbage and cauliﬂower plants. Aust. J. Soil Sci. 12, 160 –167.
Fromm, J., and Eschrich, W. (1989). Correlation of ionic movements with phloem unloading and loading in barley leaves. Plant Physiol. Biochem. 27, 577– 585.
GUOHUA XU ET AL.
Fuqua, B. D., Sims, J. L., Leggett, J. E., Benner, J. F., and Atkinson, W. O. (1976). Nitrate and chloride fertilization effects on yield and chemical composition of Burley tobacco leaves and smoke.
Can. J. Plant Sci. 56(4), 893 – 899.
Gaspar, P. E., Reeves, D. L., Schumacher, T. E., and Fixen, P. E. (1994). Oat cultivar response to potassium chloride on soils testing high in potassium. Agron. J. 86(2), 255 –258.
Gausman, H. W., Cunningham, C. E., and Struchtemeyer, R. A. (1958a). Effects of chloride and sulfate on 32P uptake by potatoes. Agron. J. 50, 90 – 91.
Gausman, H. W., Corbett, E. G., and Struchtemeyer, R. A. (1958b). Chloride deﬁciency symptoms in
potato plants. Agron. J. 50, 403.
Glass, A. D. M., and Siddiqi, M. Y. (1985). Nitrate inhibition of chloride inﬂux in barley: Implications
for a proposed chloride homeostat. J. Exp. Bot. 36(165), 556 – 566.
Goos, R. J. (1987). Chloride fertilization: The basics. Crops Soils 39, 12–13.
Gorham, J., and Bridges, J. (1995). Effects of calcium on growth and leaf ion concentrations of Gossypium hirsutum grown in saline hydroponic culture. Plant Soil 176(2), 219 –227.
Gouia, H., Ghorbal, M. H., and Touraine, B. (1994). Effects of NaCl on ﬂows of N and mineral ions
and on NOϪ
3 reduction within whole plants of salt-sensitive bean and salt-tolerant cotton. Plant
Physiol. 105(4), 1409 –1418.
Grattan, S. R., and Grieve, C. M. (1992). Mineral element acquisition and growth response of plants
grown in saline environments. Agric. Ecosyst. Environ. 38(4), 275 – 300.
Grattan, S. R., and Maas, E. V. (1985). Root control of leaf phosphorus and chlorine accumulation in
soybean under salinity stress. Agron. J. 77(6), 890 – 895.
Grifﬁth, C. J., Rea, P. A., Blumwald, E., and Poole, R. J. (1986). Mechanism of stimulation and inhibition of tonoplast Hϩ-ATPase of Beta vulgaris by chloride and nitrate. Plant Physiol. 81, 120–125.
Haeder, H. E. (1976). The inﬂuence of chloride nutrition in comparison with sulfate nutrition on assimilation and translocation of assimilates in potato plants. Landwirtsch. Forsch., Sonderh. 32(1),
Hager, A., and Helmle, M. (1981). Properties of an ATP-fueled, Cl-dependent proton pump localized
in membranes of microsomal vesicles from maize coleoptiles. Z. Naturforsch., Sect. C: Biosci.
Hajibagheri, M. A., Yeo, A. R., Flowers, T. J., and Collins, J. C. (1989). Salinity resistance in Zea mays:
Fluxes of potassium, sodium and chloride, cytoplasmic concentrations and microsomal membrane
lipids. Plant Cell Environ. 12, 753 –757.
Hang, Z. (1993). Inﬂuence of chloride on the uptake and translocation of phosphorus in potato. J. Plant
Nutr. 16(9), 1733 –1737.
Harward, M. E., Jackson, W. A., Piland, J. R., and Mason, D. D. (1956). The relationship of chloride
and sulfate ions to forms of nitrogen in the nutrition of Irish potatoes. Soil Sci. Soc. Am. Proc. 20,
Hassan, M. M., and El-Samnoudi, I. M. (1993). Effect of soil salinity on yield and leaf mineral content of date palm trees. Egypt. J. Hortic. 20(2), 315 – 322.
Hauck, R. D. (1984). “Nitrogen in Crop Production,” pp. 507– 519. Am. Soc. Agron., Madison, WI.
Hawker, J. S., and Walker, R. R. (1978). The effect of sodium chloride on the growth and fruiting of
Cabernet Sauvignon [grape] vines. Am. J. Enol. Vitic. 29(3), 172–176.
Heckman, J. R. (1995). Corn responses to chloride in maximum yield research. Agron. J. 87(3), 415 –
Heckman, J. R. (1998). Corn stalk rot suppression and grain yield response to chloride. J. Plant Nutr.
Ho, L. C., Grange, R. I., and Picken, A. J. (1987). An analysis of the accumulation of water and dry
matter in tomato fruit. Plant Cell Environ. 10, 157–162.
Hoﬁnger, M., and Bottger, M. (1979). Identiﬁcation by GC-MS of 4-chloroindolylacetic acid and its
methyl ester in immature Vicia faba broad bean seeds. Phytochemistry 18, 653 – 654.
ADVANCES IN CHLORIDE NUTRITION
Homann, P. H. (1988). Structural effects of chloride and other anions on the water oxidizing complex
of chloroplast photosystem II. Plant Physiol. 88(1), 194 –199.
Huang, Y., Rao, Y. P., and Liao, T. J. (1995). Migration of chloride in soil and plant. J. Southwest Agric.
Univ. (in Chinese) 17(3), 259 –263.
Huber, D. M., and Arny, D. C. (1985). Interaction of potassium with plant disease. In “Potassium in
Agriculture” (R. D. Munson, ed.), pp. 467– 488. Am. Soc. Agron., Crop Sci. Soc. Am., Soil Sci.
Soc. Am., Madison, WI.
Huber, D. M., and Wilhelm, N. S. (1988). The role of manganese in resistance to plant disease. In “Manganese in Soils and Plants” (R. D. Graham, R. J. Hannam, and N. C. Uren, eds.), pp. 155 –173.
Kluwer Academic Publishers, Boston.
Imas, P. (1991). Yield-transcription relationship under different nutrition conditions. M.Sc. Thesis, pp.
84–86. Hebrew University of Jerusalem, Faculty of Agriculture.
Itoh, S., and Uwano, S. (1986). Characteristics of the Cl action site in the O2 evolving reaction in PSII
particles: Electrostatic interaction with ions. Plant Cell Physiol. 27, 25 – 36.
Izawa, S., Heath, R. L., and Hind, G. (1969). The role of chloride ion in photosynthesis. III. The effect
of artiﬁcial electron donors upon electron transport. Biochim. Biophys. Acta 180, 388 – 398.
Jackson, T. L., and McBride, R. E. (1986). Yield and quality of potatoes improved with potassium and
chloride fertilization. In “Special Bulletin on Chloride and Crop Production” (T. L. Jackson, ed.),
No. 2, pp. 73–83. Potash & Phosphate Institute, Atlanta, GA.
Jacoby, B., and Rudich, B. (1980). Proton–chloride symport in barley roots. Ann. Bot. (London) [N.S.]
James, D. W., Weaver, W. H., and Reeder, R. L. (1970). Chloride uptake by potatoes and the effects of
potassium chloride, nitrogen and phosphorus fertilization. Soil Sci. 109(1), 48 – 53.
Jeong, B. R., and Lee, C. W. (1992). Growth suppression and raised tissue Cl contents in NHϩ
marigold, petunia, and salvia. J. Am. Soc. Hortic. Sci. 117(5), 762–768.
Jeschke, W. D., and Wolf, O. (1988). External potassium supply is not required for root growth in saline
conditions: Experiments with Ricinus communis L. grown in a reciprocal split-root system. J. Exp.
Bot. 39(206), 1149–1168.
Ji, G. L. (1997). Electrostatic adsorption of anions. In “Chemistry of Variable Charge Soils” (T. R. Yu,
ed.), pp. 112–139. Oxford University Press, New York.
Jing, A. S., Guo, B. C., and Zhang, X. Y. (1992). Chloride tolerance and its effects on yield and quality of crops. Chin. J. Soil Sci. 33(6), 257–259.
Johnson, C. M., Stout, P. R., Broyer, T. C., and Carlton, A. B. (1957). Comparative chloride requirements of different plant species. Plant Soil 8, 337– 353.
Johnson, G. V., and Fixen, P. E. (1990). Testing soils for sulfur, boron, molybdenum, and chlorine. In
“Soil Testing and Plant Analysis” (R. L. Westerman, ed.), 3rd ed., Vol. 3, pp. 265 –273. SSSA
Book Series, Madison, WI.
Junge, C. E. (1963). “Air Chemistry and Radioactivity,” pp. 311– 333. Academic Press, New York.
Kafkaﬁ, U. (1987). Plant nutrition under saline condition. Fertil. Agric. 95, 3 –17.
Kafkaﬁ, U., and Bernstein, N. (1996). Root growth under ionic composition stress. In “Plant Root—
The Hidden Half” (Y. Waisel, A. Eshel, and U. Kafkaﬁ, eds.), pp. 435– 452. Dekker, New York.
Kafkaﬁ, U., Bar-Yosef, B., and Hadas, A. (1978). Fertilization decision model. Soil Sci. 125, 261–
Kafkaﬁ, U., Valoras, N., and Letay, J. (1982). Chloride interaction with NOϪ
and phosphate nutrition
in tomato. J. Plant Nutr. 5(12), 1369 –1385.
Kafkaﬁ, U., Siddiqi, N. Y., Ritchie, R. J., Glass, A. D. M., and Ruth, T. J. (1992). Reduction of 13NOϪ
inﬂux and 13N translocation by tomato and melon varieties after short exposure to Ca and K chloride salts. J. Plant Nutr. 15(6 –7), 959 – 975.
Keller, J., and Bliesner, R. D. (1990). Trickle irrigation planning factors. In “Sprinkle and Trickle Irrigation” (J. Keller and R. D. Bliesner, eds.), pp. 453 – 477. Van Nostrand-Reinhold, New York.