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Alternative or Complementary Role of Foliar Supply in Mineral Nutrition

K. Mengel Justus, Liebig-University Giessen, Institute of Plant Nutrition Pohlheim, Germany Keywords: epicuticular wax, foliar spray, iron, phloem-mobility, nutrient uptake Abstract Morphology and organisation of leaf tissue is such that it accomodates the uptake of gaseous plant nutrients, whilst that of roots the uptake of water-soluble solutes. These water soluble plant nutrients are mainly supplied with fertilizers. Only in exceptional cases where nutrients are strongly fixed by soils or where aerial nutrient requirement of a crop is higher than the root uptake rates, foliar application can be adopted as a routine fertilization measure. The efficiency with which foliar applied nutrients are utilized depends on the mobility of the specific nutrient throughout the entire plant, mobility comprising long distance transport especially phloem transport as well as the symplastic transport. Potassium and nitrogen are examples of nutrients showing high mobility and when taken up by leaves they can be rapidly distributed throughout the entire plant. Calcium, sulfur, and iron show a low mobility and Ca2+ taken up by leaves cannot be transported to younger tissues or fruits where it may be required. Micronutrient requirements can generally be better met by foliar application than requirements of macronutrients because in absolute terms higher quantities of macronutrients are needed. Iron in the leaf apoplast can be mobilized by a foliar application of diluted acids. INTRODUCTION Plant nutrients are taken up both by roots and by upper plant parts, mainly leaves. During evolution specialized organs, roots and leaves, have developed which are relatively specific for the uptake of particular forms of nutrient species: gaseous nutrients such as CO2 and O2 as well as SO2 (Faller, 1968) and NH3 (Lemon and van Houtte 1980) are taken up from the atmosphere by leaves, whereas water soluble nutrients, mainly ions present in the soil solution are absorbed by roots. The morphology and anatomy of both leaves and roots are such to accommodate this difference in behaviour. Uptake of water soluble nutrients by leaves is not usally very efficient and plays no major role in plants growing in natural habitats. For crop species, however, developed by man and characterized by high yields nutrient uptake by roots may be inadequate to meet nutrient demand of the upper plant parts, especially of harvested plant parts such as fruits, grains and seeds. Here foliar application of nutrients offers a complementary means of providing nutrients during a critical phase of restricted nutrient supply. In addition it can alleviate the effects of strong binding of plant nutrients in soils as well as difficulties in the acquisition of nutrients because of particular soil conditions as can be the case for Fe on calcareous soils. Foliar application is particularly of importance for perennial fruit crops with deeprooting systems which in time may deplete the surrounding soil of available plant nutrients and where soil-surface application of fertilizers is almost without effect because of nutrient fixation as happens in K+ fixing soils. From this it follows that foliar application is not an alternative but only a complementary measure to ensure adequate nutrient supply in the nutrition of plants. THE MESOPHYLL, SYMPLASTIC AND APOPLASTIC PATHWAYS Figure 1 shows the cross section of a leaf with the various tissues including the stomata and stomata cavity. Air may easily enter the stomata within the spongy

Proc. IS on Foliar Nutrition Eds. M.Tagliavini et al. Acta Hort. 594, ISHS 2002


mesophyll and from here CO2 and O2 can penetrate the plasma membrane into the cytosol. Water soluble nutrients when sprayed on a leaf surface have to overcome considerable barries before entering the cytosol. According to more recent results also open stomata promote the uptake of foliar applied solutes (Burkhardt and Eichert, 2001). Foliar spraying, however, is frequently applied late in the afternoon or even in the evening when the air humidity is high and evaporation of sprayed droplets low. At this time of the day most stomata are closed. Under such conditions the foliar-applied solutes in many cases have to penetrate the cuticle which is a strong barrier for hydrophilic solutes (cuticle and the cuticular waxes will be discussed more in detail below). After penetration of the cuticle the solute enters the free space of the cell wall (apoplast). In leaves in contrast to roots a large proportion of this free space is filled with atmospheric gases which is several times larger than the water filled proportion (Lohaus et al., 2001). Only the water filled free space allows the diffusion of solutes and since some of this water is tightly adsorbed to cell wall structures this also restricts solute diffusion. It hence follows that only at high atmospheric humidity are fairly good diffusion conditions provided for diffusion of solutes. Besides diffusion, transport of solutes in the apoplast is brought about by mass flow with the transpiration water which moves from the xylem tissue to the leaf surfaces particulary to the stomatal cavities. This water movement may contribute to the distribution of solutes in the leaf tissue; on the other hand transpiration may lead to a considerable loss of water from the cell wall free space and hence restrict the solute diffusion. For the distribution of foliar-appplied nutrients the symplastic pathway is the more important one. Diffusion rates from the leaf surface into the free space of the cell wall depend much on the concentration difference between the nutrient concentration in the droplets sprayed on the leaf surface and the nutrient concentration in the apoplast. However, where the plasma membrane is well equipped with uptake systems (transporters and channels) for the specific nutrient its uptake provides a continuous concentration gradient driving nutrient diffusion from the leaf surface into the apoplast. This is especially the case when the uptake is not limited by low temperature and when there is a demand for the nutrient. For all those nutrients which are transported under deficiency conditions from older leaves to younger leaves , older leaves provide better foliar uptake conditions than do younger ones, since the total surface of mature leaves is larger than the total surface of the developing leaves. In addition in older leaves the cuticles are partially damaged and contain cracks which function as diffusion pathways for the solutes. Nutrient species which first become deficient in the older leaves include N, P, K, Mg, Mo, and Zn (Caballero et al., 1996) and in some species also B (Brown and Shelp, 1997). Those plant nutrients which older leaves either do not contribute or contribute little to the nutrient demand of younger leaves include Ca, S (Bouma, 1967), Mn, and Cu. These nutrients are less phloem mobile which is a great disadvantage in foliar application as will be considered more in detail below. Nutrients having once entered the cytosol can be transported from one cell to another via the plasmodesmata. In this way they can be easily distributed all over a particular tissue. They also may enter the sive cell/companion cell complex via the plasmodesmata from where they can be loaded into the phloem tissue and then are subjected to long distance phloem transport to be translocated to those tissues where they are required. This applies for the phloem mobile plant nutrients and is particularly important for N, K, Mg, Mo and Zn. In the case of N, phloem transport occurs mainly in the form of amino acids. The N of urea, ammonium and/or nitrate in foliar sprays has to be transformed into amino-N before it is subjected to long-distance phloem transport. Potassium sprayed on the leaves is quickly translocated to stems and roots as shown in an earlier work of Scharrer and Mengel (1960) some results of which are shown in Tab.1. The cytosolic concentration of Ca2+ is in the range of several M and hence extremely low. It is for this reason that the symplastic pathway actually plays no role in the distribution of Ca2+ in the tissue. Therefore the Ca2+ distribution in the leaf and especially its transport to the vascular tissue has to follow the apoplastic pathway. The


apoplast, however, comprises numerous indiffusible anions, mainly carboxylic groups of pectic and glucuronic acids, which bind Ca2+ preferentially to Mg2+, K+, and Na+ and thus restrict Ca2+ diffusion. In addition the indiffusible anions form Donnan sites (pockets) in which mainly Ca2+ is accumulated. It is for this reason that the efficiency of the apoplastic pathway for the Ca2+ transport is also poor and therefore Ca2+ which has penetrated the cuticle and entered the leaf apolast has little chance of reaching the vascular tissue and hence the systems for long distance transport. Here only the xylem transport can play a role in the distribution of Ca2+ because the sieve cell companion cell complex feeding the phloem transport system does not load the sieve cells with Ca2+. Xylem transport occurs only in acropetal direction and is of some importance for the Ca2+ supply of young fruits which still transpire in the early phase of their development. At a later stage fruits almost exclusively feed from the phloem. This is of particular importance for apples since it is clear that foliar Ca2+ supply to apple leaves is of no major importance for the supply of apple fruits with Ca2+ and it is the fruit itself which must be sprayed with a Ca2+ solution. Early experiments of Kick and Hellwig (1959) showed that foliar supply of K+ and N is very efficient if the plants are sprayed frequently while the foliar application of phosphate was inferior to the soil phosphate application. Although phosphate is phloemmobile and can be well distributed in the symplastic pathway, penetration of the cuticle by phosphates and the behaviour of phosphates in the spraying droplets on the leaf surface give rise to difficulties. EPICUTICULAR WAXES, CUTICLE AND CELL WALL STRUCTURE Figure 2 shows a simplified scheme of the layers nutrients have to traverse during their transport from the leaf surface to the plasma membrane, modified after the scheme of Franke (1967) published in a still informative review article. It shows the wax layer with the wax rodlets covering the cuticle, the primary and secondary cell wall with the plasma membrane overlaying the cytosol. The waxes mainly forming rodlets are hydrophobic and hence represent a strong barrier for water soluble substances, e.g. plant nutrients. These waxes are crystalloids which in some plant species are reinforced by polymeric aldehydes thus increasing the water repellency (Haas et al., 2001). In young leaves epicuticular waxes form an almost continuous cover and therefore water soluble solutes have virtually no chance of penetrating it. Rain drops are repulsed by this hydrophobic layer and foliar spraying solutions need the addition of a detergent (surfactant) in order to decrease the water tension of the solution so that spray droplets form a thin film on the leaf surface. The duration of this film is of importance for the quantity of nutrients taken up; the longer the contact time the higher the uptake. For this reason air humidity should be high in order to avoid a major water loss of the spraying solution because of evaporation. Along with the aging of leaves, cracks and deletions may develop in the wax layer which allow a contact of the solute with the cuticle. Environmental conditions such as acid fog (Plate I) may contribute to the deterioration of epicuticular waxes (Mengel et al., 1989) and insufficient K+ supply may affect their development (Sunxi and Rao Li-Hua, see Mengel and Kirkby, 2001). Thus as leaves age there is also an accompanying increase in the chance that foliar applied nutrients will reach the cuticle. The main components of the cuticle are cutin and wax plates which are embedded in the frame work of the cutin. Cutin is a polymeric network of C16 and C18 hydroxy fatty acids in which the carboxylic group is esterified with the hydroxyl group. These esterified sites are hydrophilic (-CO-O-) and therefore allow the diffusion of hydrophilic solutes across the cutin layer and hence their contact with the primary cell wall. This is made up of a cellulose framework cross-linking glycans which are embedded in a matrix of pectic polysaccharides. In addition primary cell walls contain proteins. The building strutures of the primary cell wall are mainly hydrophilic. The pectic matrix provides the porosity of the primary cell wall with spaces of a diameter in the range of < 5 nm. Considering that sucrose, Ca2+, and K+ have a diameter of 1.0, 0.82, and 0.66 nm, respectively, it is obvious that the pectic matrix of the primary cell wall represents a barrier for large


molecules, e.g. for chelated plant nutrients. In addition, the pectic matrix carries in indiffusible anions which may form Donnan sites capable of trapping cations, especially Ca2+. Such Donnan sites may also be formed by borate esters bridging two rhamnogalacturonan strands as shown in Fig. 3 (Carpita et al., 1996). The secondary cell wall is formed on the plasma membrane side of the primary wall (Fig. 2). The main framework of the secondary wall consists of cellulose in which other compounds like lignin and even hydrophobic substances can be embedded, such as suberin in roots. In most cases the secondary cell wall in leaf tissue represents no major barrier for water-soluble solutes. PROPERTIES AND EFFICIENCY OF FOLIAR APPLIED NUTRIENTS The efficiency with which foliar applied nutrients are utilized depends much on the total demand of the particular nutrient. This is much higher for macronutrients than for micronutrients since the quantity of foliar applied nutrients taken up is low in absolute terms and more closely follows micronutrient than macronutrient demand. Of the macronutients it is foliar N application which especially plays a major role in practical crop production, e.g. the post harvest foliar nitrogen application to fruit trees. Tagliavini et al. (1998 ) reported that peach leaves were able to take up before leaf senescence appreciable amounts of urea N and 60 to 70% of the urea N intercepted by the canopy was translocated before leaf abscission into the wood , mainly the roots and contributed to the N supply of trees in the following spring. Basically the N carriers can be foliar applied in the form of urea, nitrate, and ammonium. Urea is a non-electrically charged molecule which is water soluble but has also some hydrophobic properties. For this reason it is particularly suitable for foliar application as it can relatively easily penetrate the epicuticular wax as well as the cutin layer. Urea is frequently sprayed together with pesticides (Sturm et al., 1994). As yet not much is known about the uptake mechanism of urea by the plasmalemma. According to earlier results of Kirkby and Mengel (1970) obtained with roots, uptake seems to take place via diffusion across the plasmalemma and thus be dependent on the urea concentration in the nutrient solution. Since urea possesses has some hydrophobic properties it is feasible that it penetrates the plasma membrane without the involvement of a particular transporter system. Urea thus entering the cytosol could quickly be hydrolyzed by urease which is present in most plant cells (Freney et al. 1992) and the resulting NH3 is metabolized in the GOGAT pathway. By these processes a urea concentration gradient between the urea on the leaf surface and on the plasma membrane surface could be maintained to drive the urea uptake by leaves. Urea leaf application is common for arable crops (Sturm et al. 1994) and even more for perennial fruit crops. After fruit maturation, in leaves of perennial fruit crops including grapevine proteolysis dominates and the resulting glutamine is required for the synthesis of arginine (Prior, 1997) which is stored in the wood and mobilized for leaf growth in spring. Since NH3 produced by urea hydrolysis can be quickly used for the synthesis of glutamine in only one step (glutamine synthetase) post harvest foliar application should be particularly efficient in promoting shoot and leaf formation in spring. Most nitrates are well soluble in water and thus appropriate for foliar application. The accompanying cation may play a role in nitrate diffusion across the cuticle since as a consequence of their electrical charges cation and anion have to move together. Ca2+ as accompanying cation may be bound to the indiffusible cell wall anions and thus restricting the nitrate transport. Ammonium, K+, and Na+ are more appropriate accompanying cations since they remain virtually unbound to cell wall structures. Nitrate uptake rates depend on the efficiency of nitrate transporters in the plasma membrane of leaves. Kosegarten et al. (1999) found that the apoplastic pH of sunflower leaves was significantly raised when the plants were fed with nitrate as compared with ammonium. This is evidence that in leaves a H+/nitrate transporter functions in the transport of nitrate across the plasma membrane analogous to the nitrate transporter in roots. There is not much known about NH4+ transporters in leaf plasma membranes. The


results of Kosegarten et al. (1999) are not consistent with a H+/NH4+ cotransport across the plasma membrane in leaves as was found for roots by Schachtman and Schroeder (1994). Such an uptake system in leaves is not likely since NH4+ taken up by roots in contrast with nitrate is immediately metabolized in situ for amino acid synthesis and not transported into the upper plant parts (Kosegarten and English, 1994). Theroretically NH4+ may penetrate the leaf plasma membrane by facilitated diffusion a process driven to a large extent by the strong electrochemical potential difference for NH4+ between the leaf apoplast and the cytosol. Such a transport requires specific channels and it is not yet known whether these exist in leaf plasma membranes as they are known to occur in the plasma membranes of roots (Ullrich, 1992). Of the various K+ salts it is frequently the K nitrate which is foliar applied. This compound is highly soluble in water and both ion species diffuse relatively easily through the cuticle and the free space of the cell wall. Both are taken up at high rates and hence provide a concentration gradient great enough to drive the ions from the leaf surface to the surface of the plasma membrane. As already mentioned above, K+ foliar application may be useful for deep- rooting perennials grown on K+ fixing soils. It may also have a beneficial effect on annual crops as is shown for cotton in Tab.2 (Howard et al., 1998). In this experiment there is an effect of the H+ buffer solution sprayed together with the K salt on the leaves. Only when the buffer solution had a pH of 4 there was a significant yield increase obtained regardless of whether K+ was applied as nitrate or sulfate. The interpretation of this result is difficult but it seems that the H+ buffered foliar applied solution had an impact on the apoplastic pH, the low pH presumably favouring the K+ uptake. Such a behaviour is in accord with the assumption that the K+ uptake takes place via a K+/H+ symport across the plasmalemma as it was found for roots by Schachtman and Schroeder (1994). These are high affinity K+ transporters which means that they deplete the K+ concentration to a very low level which in turn would favour the K+ diffusion from the leaf surface to the plasma membrane. Most Mg salts are soluble in water and their ions may diffuse from the leaf surface across the cuticle into the free space of cell walls. There is hardly anything known about Mg2+ transporters in the plasma membrane neither of roots nor leaves. In the root medium Mg2+ uptake can be severely restricted by K+ (Schimansky, 1981). The early work of Mulder (1950) reported that high levels of available K+ in soils induced Mg deficiency symptoms in apple trees. Such a trait is consistent with the assumption that both cation species compete for the negative charge of the cytosol and that the uptake is mediated through specific cation channels. Such a competition may restrict the Mg2+ uptake of roots considerably, but in the leaf apoplast presumably high K+ concentrations do not prevail to restrict the uptake of foliar applied Mg2+ and under such conditions Mg foliar application will be beneficial. Sulfate transport from older to younger leaves is much restricted and from this behavour may be concluded that sulfate sprayed on older leaves will be unlikely to reach reach the younger plant parts where S deficiency may occur. The poor mobility of Ca2+ in plant tissues has already been considered above. The mobility of Mn2+ in the leaf apoplast may be similar to that of Ca2+. In contrast to Ca2+, however, Mn foliar application can cure Mn deficiency. For Ca2+ it is mainly the fruits that are deficient (apples, tomatoes) whereas for Mn deficiecny symptoms appear on the leaves and this reason Mn foliar application is an efficient measure to alleviate the deficiency. This is particularly the case in alkaline organic soils in which Mn2+ is quickly oxidized to MnIII or MnIV which are not available. Draycott and Farley (1973) in comparing soil application of Mn silicates and Mn oxides with Mn foliar sprays found that soil application did not prevent Mn deficiency in sugar beet whereas Mn foliar application corrected the deficiency and increased sugar yields. According to Ozaki (1955) spraying a MnSO4 solution is more efficient than Mn chelates in curing the deficiency. Zinc deficiency is frequently found on calcareous soils (Cakmak et al., 1997) because its availablity decreases with soil pH. Zn2+ is specifically adsorbed and even


included by soil carbonates (Donner and Lynch, 1989). Field trials carried out in Egypt (Serry et al., 1974) showed that on the alluvial soil with a moderate deficiency Zn foliar application was as efficient as the soil application. On the calcareous soils where deficiency was more pronounced the foliar application resulted only in a modest yield increase whereas a soil application of 19 kg Zn/ha produced a marked increase in grain yield. For annual field crops soil application of Zn is generally superior to foliar spray. This may not be applicable to perennial fruit trees with deeper rooting systems where the soil application of Zn fertilizer even when incorporated into the upper soil layer of a calcareous soil will have no major effect because of Zn fixation in the deeper soil layers. Here the foliar application is the appropriate measure. On acid organic soils (Oxisols) Zn may be complexed by the organic matter resulting in Zn deficiency in citrus and coffee grown in Brasil (Rosolem and Sacramento, 2001). Foliar application of Zn- lignosulfate, Zn EDTA, and ZnSO4 with rates of about 1.7 kg Zn/ha, all were capable of increasing the Zn concentration in the leaves to an adequate level. In one of four field trials ZnSO4 was superior to the chelated Zn forms. Copper is very strongly bound in acidic organic soils (Beringer, 1963). According to McBride 1989) Cu2+ can react with carboxylic groups as shown below. R-COO- + Cu2+ ! R-COOCu+ Such Cu2+ bindings may also occur at the indiffusible anions in cell walls and hence severely restrict the Cu2+ diffusion to the uptake sites in the plasma membrane. More than forty years ago Keller and Deuel (1958) reported that Cu2+ is strongly bound in the apoplast. Also leaf scorching frequently found after foliar application of CuSO4 may result from this binding. Reith (1968) carrying out field trials on Cu deficient glacial sandy soils obtained a significant grain yield increase with CuSO4 soil application with rates of 1 to 18 kg Cu/ha. Foliar Cu application was not as efficient as soil application although foliar application gave higher Cu concentrations in straw and grains as shown in Tab. 3. This is an interesting finding . Obviously the Cu2+ applied by foliar application remained mainly in the apoplast and was translocated to the grains by transpiration. The soil applied Cu2+ taken up by roots was transported to sites of physiological activity where it was required and although the concentration in the grains was only slightly increased by Cu, soil application gave a marked grain yield increase. In the straw Cu soil application did not raise the Cu concentration; total Cu uptake, however, was increased because of a substantial increase in straw yield. This example shows that foliar application of Cu may be inefficient. Boron in the form of boric acid is a non-charged molecule under the pH conditions of the apoplast. For this reason it is relatively mobile in the apoplast and frequently used in foliar application. Its main physiological target is the primary cell wall of meristematic tissues where it is essential for the construction of the pectic framework (see Fig. 3). Reproductive growth may exceed the vegetative growth B requirement of (Nyomara, 1997). Hence foliar B application at flowering is beneficial. Such positive effects are particularly obtained in plant species in which B is phloem-mobile. According to Brown and Shelp (1997) there are plant species e.g. apple trees, in which B is quickly distributed over the entire plant whereas in others this is not so, e.g. walnut trees. High mobility means that B foliar application is much effective. The impact of soil and foliar B application on citrus fruit growth and quality is shown in Tab.4. (Foroughi et al., 1973). Boron supply promoted fruit size and decreased the peel thickness. From the data it is obvious that foliar application was superior to soil application. On heavy soils with a high pH, B(OH)4- is strongly adsorbed by clay minerals. In such cases foliar application is the appropriate measure. Absolute Fe deficiency is very rare and plays virtually no role in crop production. It may appear in sterile soils which lack the microorganisms for the synthesis of siderophores required for Fe transport from the soil medium to plant roots (Masalha et al., 2000). Iron deficiency occurs on alkaline and calcareous soils where the high HCO3concentration in the soil solution results in a high pH in the root apoplast which restricts


the reduction of FeIII chelates and hence the uptake of Fe2+ into the cytosol (Kosegarten and Koyro 2001). In such cases the Fe concentration in the root apoplast is high (Kosegarten et al., 1998, Kosegarten and Koyro, 2001). Fe-chelates are used for soil and foliar application their effect depending much on the time of application as discussed by Tagliavini and Rombol (2001) in a useful review article. Iron is only to a very low degree phloem-mobile and therefore older leaves do not contribute to the Fe supply of younger ones as is also the case of Fe sprayed on older leaves. Iron is unique in the plant nutrients in that its concentration in the plant particularly in leaves is not related to its physiological Fe status and frequently leaves may contain high Fe concentrations but still are chlorotic (Mengel et al., 1984, Rmheld, 2000). Iron chlorosis in leaves is induced by nitrate nutrition (Kosegarten et al., 1999) because nitrate nutrition leads to a pH increase in the leaf apoplast due to the NO3-/H+ cotransport. This pH increase restricts the activity of the FeIII reductase and therefore the uptake of Fe2+ into the cytosol (Kosegarten et al. 1999). In recent investigations Kosegarten et al. (2001) depressed the apoplastic leaf pH from 5.4 to 4.9 by spraying citric acids on chlorotic leaves of sunflower. This pH decrease resulted in a regreening of leaves without a change of their Fe concentration as shown in Tab.5. The authors interprete their findings in terms of the activation of the plasmalemma FeIII reductase at the low pH followed by the uptake of Fe2+ from the apoplast into the cytosol. This interpretation is consistent with results of Sahu et al. (1987) who cured Fe chlorosis in peas by spraying the leaves with a diluted sulfuric acid. In their experiment the foliar application of sulfuric acid was more efficient in curing the chlorosis and increasing pod yield than the foliar application of a Fe-EDDHA. It should be emphasized that acidic sprays are only efficient if there is enough Fe trapped in the apoplast. This is in accord with foliar application to kiwifruit by Tagliavini et al (1995) who found that spraying of Fe-DTPA as well as a foliar application of citric acid resulted in leaf regreening; FeDTPA, however, was more efficient than citric acid. Field trials caried out with kiwifruit, peaches, and pears in various mediterranean countries provide evidence that fruit yield is positively correlated with the leaf chlorophyll concentration which was increased by foliar spraying of Fe-DTPA, FeSO4 + citric acid and FeSO4 alone. In FeSO4 the Fe is divalent and in this form can directly traverse the plasmalemma and hence should be well available for other physiological processes (Fox and Guerinot, 1998). It should be born in mind, however, that in the presence of O2 ferrous Fe can be rapidly oxidised to ferric Fe. In the Fe chelates used for foliar application the Fe is trivalent and therefore must be reduced before taken up into the cytosol. This reduction may be restricted if the apoplastic pH is high, a situation which may occur if the plants are exclusively fed with nitrate-N. Finally it should be stressed that the long-distance transport of Fe is not efficient which means that the Fe translocation from a sprayed leaf to young meristematic tissues where the Fe is required for developing new leaves (Kosegarten et al., 1998) is much retricted. These are the reasons why Fe foliar application frequently is not as effective as expected and that foliar applications must often be repeated in order to obtain a satisfactory effect. CONCLUSION The efficiency of foliar applied nutrients depends much on their mobility in the entire plant. Plant nutrients which are phloem- mobile may easily be transported from the leaf on which they were sprayed to any other plant organ where they are required. For such nutrients as is the case for nitrogen and K+ the whole foliage may function as acceptor. This does not apply to Ca2+, sulfate and Fe for which the phloem transport is of no major importance and foliar application of these nutrients is frequently inefficient. In foliar applied FeIII- chelates the ferric Fe must be reduced before it can be taken up into the cytosol. Reduction can be restricted by a high apoplastic pH induced by nitrate uptake via H+/nitrate cotransport. Foliar plasmalemma uptake systems are of importance for the diffusion of sprayed nutrients from the leaf surface to the plasma membrane. Not much is known about such uptake systems in the leaf apoplast. This is of particular interest for urea which is often used in foliar application. The leaf apoplast contains numerous


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Table 2. Effect of H+ buffering the foliar spray solution on lint yield of cotton. Yield data are the average of four years (Howard et al., 1998). K Salt Solution pH Lint Yield (kg/ha) Control, no foliar applic. 1185dc KNO3 9.3 1195de 6.0 1222cd KNO3 KNO3 4.0 1291ab K2SO4 9.5 1190de K2SO4 6.0 1196e 4.0 1340a* K2SO4 Yield data with the same letter do not differ significantly for P = 0.05.


Table 3. Effect of Cu soil and foliar application to spring barley and oats on grain and straw yield and Cu concentrations in grain and straw. Means of six field trials (Reith, 1968). Soil application: 3.7 kg Cu/ha, foliar application: 180 g/ha. Control, no Cu. 3500 4900 1.9 1.6 Soil appl. 4750 5600 2.0 1.6 Foliar appl. 4400 4625 3.8 2.7

Grain yield, kg/ha Straw yield, kg/ha Cu concentr., g/g DM grain Cu concentr., g/g DM straw

Table 4. Effect of soil and foliar B application on fruit diameter and peel thickness of Citrus aurantium. Borax was used for soil application and Solobur for foliar application (Foroughi et al., 1973). Treatment Control, no B 50 g B/tree, soil appl. 100 g B/tree, soil appl. 200 g B/tree, soil appl. 15 g B/tree, foliar appl. 30 g B/tree, foliar appl. 60 g B/tree, foliar appl. LSD, P> 0.01 5.9 3.1. Fruit , mm 48.5 62.0 62.0 63.5 62.0 62.4 68.0 Peel thickn., mm 25.8 18.0 18.5 16.5 15.2 15.0 14.8

Table 5. Effect of citric acid foliar application on Fe concentration and chlorophyll concentration in leaves and on leaf apoplastic pH (Kosegarten et al., 2001). Treatment Control, no spray Citric acid spray *** significant at <0.001. g Fe/ g DM 25.2 23.2 mg Chlorophyll/g FM 447 889*** Apoplastic pH 5.4 4.9



Fig. 1. Cross section through a leaf.

Fig. 2. Simplified scheme of the outer wall of an epidermal leaf cell. Modified after Franke (1967).


Fig. 3. Upper part: Boric acid forming with cis diol a borate ester. Lower part: Diester bonds of anionic borate bridging two strands of rhamnogalacturonan II, a constituent of the primary cell wall.


Plate I. Scanning electron micrographs of the structure of epicuticular waxes in the stomata area of Norway spruce nedles (Picea abies) of the current year (Mengel et al. 1989). A Control, application of fog with pH 5.0 B Application of acid fog with pH 3.0