Background

Micronutrients play a pivotal role in various metabolic functions through enzyme activation, chlorophyll synthesis and nutrient regulation in plants (Tripathy et al., 2015). Genotypic variation in nutrient absorption is amply demonstrated in crop plants (Clark, 1983). Nutrient absorption was also found to be influenced by rootstocks used (Cook and Lider, 1964; Downton, 1977) and varieties grafted on a same rootstock (Shikhamany et al., 2018). While organic acid metabolism was found to direct the ion absorption by plants (Jacobson and Ordin, 1954), metabolism was related to nutrient ion exchange in plant tissues (Barber and Russell, 1961). Interactions between nutrients in the foliar tissues are well documented (Fageria, 2001). They occur when the supply of one nutrient enhances or reduces the absorption and utilization of other nutrients. Interaction of major nutrients with micronutrients was found to vary with stionic combinations in grape (Shikhamany et al., 2018). These investigations were carried out to assess the interaction of micronutrient status with major nutrient absorption, and identify those micronutrients which have positive relationship with major nutrient absorption, and the stionic combinations in which such relationship exists. Identification of micronutrients that have synergism with any major nutrient or antagonism with sodium, particularly in saline alkali soils, in a given stionic combination will help in their management. Foliar absorption of micronutrient is the added advantage in the regulation of their status in vines.

1 Materials and Methods

Soil and petiole samples were collected from 38 vineyards each of Thompson Seedless on own root, Thompson Seedless on Dog Ridge rootstock, 2A clone on own root and 2A clone on Dog Ridge in Nashik and Sangli districts of Maharashtra during 2015-16 cropping season. All the vineyards were in the age group of 4-6 years and received varying levels of nutrients. The soils of the vineyards surveyed belonged to the order ‘Vertisols’ with pH in the range of 7.76±0.52 and EC 0.604±0.428 dSm^-1. All the vines selected for the study were planted at 2.7 x 1.8 m, trained to extended Y trellis and pruned to have 30±2 canes/vine. One hundred petioles of leaves opposite to flower clusters were collected at full bloom from each vineyard and soil samples from 15-30 cm depth at 60 cm away from the vine stem at back pruning before the application of fertilizers. Available contents of N, P, K, Ca, Mg, S and Na in soil and their contents, in addition to Fe, Mn, Zn and Cu in petioles were determined by standard analytical methods suggested by the AOAC. Absorption of N, P, K, Ca, Mg, S and Na was derived by the ratio of petiole contents to their respective soil content to normalize the variation in petiole contents due to variation in the applied nutrients and the available nutrients in the soil. Linear, quadratic and multiple regression equations were fitted to elucidate the relationship of petiole contents of Fe, Mn, Zn and Cu with the absorption of N, P, K, Ca, Mg, S and Na.

2 Results and Discussion

A perusal of the ranges, means and coefficient of variation in major nutrient absorption by Thompson seedless and 2A clone on their own root and Dog Ridge rootstock (Table 1) would reveal that a considerable variation existed in the absorption. This variation could be due to the affinity of roots for specific nutrient ion/ions (Cook and Lider, 1964; Downton, 1977), physiological demand by the scion variety (Jacobson and Ordin, 1954; Barber and Russell, 1961), interaction among majior nutrients (Bergman et al., 1960; Epstein, 1972) and the interaction with micronutrients (Shikhamany et al., 2018). Variation in the absorption of individual major nutrient due to micronutrient status of the vines is presented in the foregoing pages.

2.1 Variation in the absorption of nitrogen

Nitrogen absorption was not influenced significantly by any of the micronutrients in any of the stionic combinations (Table 2). Multiple regression analysis revealed that Fe, Mn, Zn and Cu together influenced the N absorption significantly in Thompson Seedless on its own root (TS/OR) and 2A clone on Dog Ridge rootstock (2A/DR). While they accounted for 10.8 per cent variation in the absorption in TS/OR, 17.3 per cent in 2A/DR. Major positive contribution was by Cu in the former combination, whereas by Zn and Cu in the latter (Table 3).

2.2 Variation in the absorption of phosphorus

Correlations of micronutrient status of vines with their absorption of major nutrients revealed that phosphorus absorption was influenced positively by Mn in TS/DR and 2A/OR and Cu in TS/DR but negatively by Zn in TS/DR (Table 2). The real positive effect of Zn in TS/DR was masked by the inherent less capacity of the stionic combination in P absorption (Table 3), resulting in the negative correlation (Table 2). The positive effect of Zn in TS/OR was masked by the negative effect of other micronutrients and the negative effect of Zn by the positive effect of Mn in 2A/OR (Table 3), resulting in non-significant correlation of Zn with P absorption (Table 2). 

All micronutrients together influenced P absorption significantly in TS/OR, TS/DR and 2A/OR. They determined the P absorption by 38.9 per cent TS/DR, while by 24.7 and 23.2 per cent respectively in TS/OR and 2A/OR. Zinc contributed positively most, towards the determination of P absorption in TS/OR and TS/DR. Whereas, it contributed negatively, while Mn did positively in 2A/OR (Table 2). Difference in the contribution of Zn and Mn could be attributed to the preferential absorption of P, Zn and Mn in different stionic combinations (Kalbhor et al., 2017). Plant genotypes differ in the uptake, translocation, accumulation and use of nutrient elements (Clark, 1983) and P interferes with Zn at the level of plant metabolism involving its uptake, translocation and utilization (Haldar and Mandal, 1981).

Quadratic function explained better the variation in P absorption due to Mn in TS/DR and 2A/OR; Zn in TS/DR and 2A/DR, and Cu in TS/DR (Table 4). Increasing levels of petiole Mn were associated with higher absorption of P in TS/DR and 2A/OR with threshold levels of 66 and 129.8 ppm respectively (Table 5), beyond which the P absorption increased steadily (Figure 2). Response of P absorption to petiole levels beyond 200 ppm was more in 2A/OR than in TS/DR. Response of P absorption to petiole Zn was different in TS/DR and 2A/DR. While it was positive in 2A/DR, negative in TS/DR. Optimum level of petiole Zn beyond which the p absorption reduced was 121.1 ppm in TS/DR, while the same content was the threshold level beyond which P absorption increased in 2A/DR (Figure 3). This discrepancy can be attributed only to 2A, the scion variety, but not to Dog Ridge rootstock, which was common for 2A and Thompson Seedless. Increasing levels of Mn were found to increase the concentration of P in rice plants (Lidon, 1999). Synergism between Mn and P was attributed to the acidifying effect of P and also to the antagonism of Mn with Zn (Haldar and Mandal, 1981), which in turn is antagonistic to P. The negative relationship between P and Zn is well established in field crops (Fageria, 2001) and observed in grape too (Shikhamany et al., 2018). Lack of synergism between P and Mn and antagonism of P with Zn in other stionic combination could be due to differential absorption of nutrients by different stionic combinations of grape (Kalbhor et al., 2017). Increasing levels of petiole Cu were associated with increased P absorption in TS/DR (Figure 4) with threshold level of 111.9 ppm (Table 5). Copper content of petioles correlated positively with P absorption in TS/DR. A perusal of Mulder’s chart would reveal antagonism between these nutrients. This discrepancy could be due to differential sprays of copper fungicides.

Since Phosphorus status of vines is implicated in fruit bud formation (Srinivasan and Mullins, 1981), foliar feeding of Mn and Cu could be beneficial but not Zn in the absorption of available soil P in TS/DR. This advantage can be realized by Mn sprays in 2A/OR but Zn in TS/DR.

2.3 Variation in the absorption of potassium

Petiole contents of Mn and Cu in TS/DR and Cu in 2A/DR correlated positively with potassium absorption, while that of Mn negatively in 2A/OR and 2A/DR (Table 2). Multiple regression analysis revealed that the four micronutrients together determined the K absorption more on Dog Ridge rootstock compared to own rooted varieties (Table 3). Maximum positive contribution was by Cu in 2A/DR, but most negatively in TS/DR. The negative effect of Cu was nullified by the positive effects of Mn and Zn in the latter stionic combination (Table 3) resulting in positive correlation of petiole Mn with K absorption but non-significant correlation with Zn (Table 2). The positive contribution of Zn in K absorption as masked by the negative contribution of Cu on own rooted vines (Table 3) resulting in non-significant correlations (Table 2).

Quadratic function was found to be the better fit than linear function to explain the variation in K absorption due to Mn and Cu in vines on Dog Ridge rootstock and Mn alone in 2A/OR (Table 4). Increasing levels of petiole Mn were associated with higher absorption of K in TS/DR, optimum being 435.9 ppm above which the K absorption reduced. Contrarily they were associated with reduced absorption in 2A/OR up to a threshold level of 210.2 ppm above which the absorption increased (Table 5). Increasing levels of petiole Mn were found to reduce K absorption in TS/OR and 2A/DR (Figure 2). Differential effect of Mn on K absorption in different stionic combinations can be attributed to the genotypic differences as discussed earlier. Absorption of K also increased at higher rates above the threshold levels of 105.4 and 155.6 ppm of Cu respectively in TS/DR and 2A/DR (Table 5). Response of K absorption to increasing levels of petiole Cu was relatively nil in vines on their own root. This difference could be due to differential sprays of copper fungicides and preferential absorption of K by Dog Ridge roots (Kalbhor et al., 2017).

Potassium being a very important nutrient for grapes in determining the cluster size, shoot maturity and quality through its crucial role in enzyme activity and metabolism, its absorption can be increased by foliar application of Mn and Cu in Thompson Seedless on Dog Ridge rootstock, Mn in 2A clone on its own root and Cu in 2A on Dog Ridge.

2.4 Variation in the absorption of calcium

Correlations of micronutrient status of vines with the absorption of calcium revealed positive relationship of Ca with Fe and Mn respectively in TS/OR and 2A/DR (Table 2). Petiole contents of Fe, Mn, Zn and Cu together could account for 27 per cent variation in Ca absorption in TS/OR and 21 per cent in 2A/DR. While the predominant contribution of Zn was positive in TS/OR, it was negative in 2A/DR (Table 3). Highest contribution towards the determination of Ca absorption was of Fe in 2A/OR. The positive effect of Mn in TS/OR was negated by Cu, of Fe by Zn in 2A/OR, while the negative effect of Zn was offset by Mn in 2A/DR (Table 3), resulting in non-significant correlations (Table 2). Quadratic function was found to explain better the variation in Ca absorption due to Fe and Mn in TS/OR and 2A/DR, and Fe alone in 2A/OR (Table 4). Increasing levels of petiole Fe were associated with increased absorption of Ca in all stionic combinations except TS/DR, in which the rate of reduction in Ca absorption was very low (Figure 1). The rate of increase in absorption was higher after the threshold levels of 116.7 and 100 ppm of Fe respectively in TS/OR and 2A/OR. In 2A/DR, the optimum level of Fe was 466.7 ppm, beyond which the increasing levels of petiole Fe was associated with reduced absorption (Figure 1). Antagonism between Ca and Fe resulting in lime induced iron chlorosis is a universal phenomenon in many field and horticultural crops, but the synergistic effect of Fe on Ca absorption reveals that the reciprocal relationship is not a rule. This could be attributed to foliar application of Fe and the high levels of available soil calcium in the vineyards under study.

Calcium absorption also increased with increasing levels of petiole Mn in TS/OR and 2A/DR, but reduced in TS/DR and 2A/OR (Figure 2). It increased at higher rates beyond the threshold level of 106.3 ppm of Mn in TS/OR, whereas increased up to the optimum level of 500 ppm beyond which decreased in 2A/DR (Table 5). Differential absorption was attributable to the differential preference for the absorption of Ca and Mn by different stionic combinations as explained earlier.

The results indicated that foliar feeding of Mn could be a potential tool to increase the absorption of Ca in Thompson Seedless vines on their own root and 2A clone on Dog Ridge rootstock. 

2.5 Variation in the absorption of magnesium

Petiole contents of Zn in TS/OR and Mn in 2A/OR correlated positively with magnesium absorption (Table 2). Multiple regression analysis revealed that the four micronutrients together determined the Mg absorption most (26.5 per cent) followed by 2A/DR (23.2 per cent), least being in TS/OR (15.9 per cent). The negative contribution of Zn was highest in 2A/OR and 2A/DR, while the contribution of Zn was positive and highest in the determination of Mg in TS/OR. Contribution of Cu was highest and positive in vines on Dog Ridge rootstock (Table 3). Negative contribution of Zn was nullified by the positive contribution of Mn in 2A/OR, resulting in its positive correlation with Mg absorption (Table 2).

Quadratic function was found to be the better fit than linear function to explain the variation in Mg absorption due to Mn in 2A clone vines on Dog Rodge rootstock, and Zn in Thompson Seedless on their own root, and in 2A clone either on its own root or Dog Ridge rootstock, but linear function was better in 2A on its own root (Table 5). Increasing levels of petiole Mn were associated with increased absorption of Mg in 2A on own root as well as on Dog Ridge rootstock. While Mg absorption increased up to 650 ppm of petiole Mn in 2A/DR, the response was linear in 2A/OR (Figure 2). Magnesium absorption increased at higher rates beyond the threshold levels of 65.2 and 112.9 ppm of Zn respectively in TS/OR and 2A/DR, but reduced drastically beyond 96.7 ppm in 2A/OR (Table 5; Figure 3).

Although Mg status of vines can be increased by its foliar application, it can also be increased by foliar sprays of Mn and Zn in 2A clone on Dog Ridge rootstock, Mn sprays in own rooted 2A and Zn sprays in own rooted Thompson Seedless.

2.6 Variation in the absorption of sulphur

Correlations of petiole micronutrient contents with the absorption of sulphur absorption revealed that S absorption was influenced positively by Mn and Cu in TS/DR and by Fe in 2A/OR, but negatively by Zn in 2A/DR (Table 2). All micronutrients together influenced S absorption significantly in TS/DR and 2A/OR. They determined the P absorption by 38.1 and 18.1 per cent respectively in TS/DR and 2A/OR. Positive contribution of Mn was highest in TS/DR, while that of Fe in 2A/OR. Petiole Mn alone accounted for 32.8 per cent variation in sulphur absorption, while Zn and Cu respectively for 20 and 19 per cent in TS/DR in the quadratic function. Fe alone accounted for 13.4 per cent in 2A/OR (Table 4). Quadratic relationship was found better to explain the variation in absorption of S due to Fe in 2A/OR, Zn and Cu in TS/DR, but linear relationship was a good fit for Mn in TS/DR (Table 5). 

Increasing levels of petiole Mn were associated with increase in S absorption linearly (Figure 2), and increased at higher rates above the threshold level of 149.3 ppm of Cu (Figure 4), but decreased above 118.8 ppm of Zn (Figure 3). Sulphur absorption increased with increasing levels of Fe up to 307.6 ppm in 2A/OR (Table 5; Figure 1). Positive effect of sulphur application on the uptake of Fe, Mn and Zn was observed and attributed to the reduced soil pH, which is conducive for their uptake (Soliman et al., 1992), but the information on the effect of micronutrient status of vines on S absorption is not available.

Sulphur deficiency in grapes is not common. However, in calcareous soils, where its availability is impaired, foliar application of Fe in 2A clone on its own root, and Mn, Zn and Cu sprays in TS/DR, if their petiole contents are below their respective optimum /threshold levels.

2.7 Variation in the absorption of sodium

Sodium is not recognized as an essential nutrient element, but it can substitute K in its physiological functions in grape (Shikhamany and Sharma, 2008). Since it is adsorbed relatively more easily on to clay surface than K, it is a strong antagonizer of K and can result in sodium toxicity. Petiole Mn contents in TS/DR and 2A/OR, and Zn contents in 2A/DR were associated positively with sodium absorption, while those of Fe and Zn in 2A/OR (Table 2). Multiple regression analysis indicated that, all the micronutrients studied, together determined the absorption of Na by 45.2 per cent in 2A/OR. Petiole Fe and Mn contents contributed most towards the determination. The determination of absorption was 23.6, 16.8 and 15.4 per cent respectively in TS/OR, TS/DR and 2A/DR. While Zn contributed most, negatively in TS/OR and TS/DR, it contributed positively in 2A/DR. Petiole Cu also contributed positively in 2A/DR (Table 3). The quadratic regression analysis revealed that, Mn alone accounted for 39.1, 31.9 and 10.7 per cent variation in Na absorption respectively in 2A/DR, 2A/OR and TS/OR, While Zn did so for 22.9 and 17.4 per cent, and Fe for 20.1 and 28.9 per cent respectively in 2A/OR and 2A/DR (Table 4). Higher contribution of Fe, Mn and Zn individually in 2A/OR and 2A/DR than the combined contribution by all the micronutrients is the indication of their mutual antagonism.

Increasing levels of petiole Fe was associated with increasing absorption of Na (Figure 1) with the optimum level of 516.6 ppm in 2A/DR (Table 5), while with reduced absorption in 2A/OR (Figure 1) beyond 63.6 ppm of petiole Fe (Table 5). Increasing levels of petiole Mn above 209.6, 257.1 and 380.8 ppm respectively in TS/OR, 2A/OR and 2A/DR were associated with reducing absorption of Na (Table 5; Figure 2). Increasing levels of petiole Zn above 117.1 ppm were associated with reduced absorption, while with increasing absorption above 66.7 ppm in 2A/DR. 

Hence, foliar sprays of Mn to increase its petiole content to more than 209.6 ppm in TS/OR, sprays of Fe, Mn and Zn to increase their levels respectively to more than 63.6, 257.1 and 117.1 ppm in 2A/OR, and to more than 516.6, 380.8 and 66.7 ppm respectively of Fe, Mn and Zn contents in 2A/DR could help reduce the absorption of Na.

Relationship of micronutrients with the absorption of major nutrients varied with their levels in petioles and stionic combinations. Relationship of Fe at levels below 100 ppm was negative with Ca absorption in TS/OR and 2A/OR, but positive in 2A/DR, The relationship got reversed above 466.7 ppm. Levels above 63.6 ppm of Fe were related negatively to Na absorption in 2A/OR but positively in 2A/DR. Mn levels below 210 ppm varied negatively with K absorption in 2A/OR but positively in TS/DR and the relationship reversed above 435.9 ppm of Mn. So also levels below 106.3 ppm were associated negatively with Ca absorption in TS/OR but positively in 2A/DR and the relationship reversed above 500 ppm of Mn. Relationship of petiole Mn with Na absorption was negative in TS/OR but positive in 2A/OR and 2A/DR at levels above 209.6 ppm, whereas at levels above 257.1 ppm the relationship turned negative in 2A/OR and so in 2A/DR above 380.8 ppm. While Zn levels above 121 ppm had negative relationship with P absorption in TS/DR, they had positive relationship in 2A/DR. Similarly Zn was associated negatively with Mg absorption in TS/OR at petiole levels below 65.2 ppm, but positively in 2A/OR. Petiole Zn levels up to 96.7 ppm were associated positively with Mg absorption in 2A/OR but negatively in 2A/DR. Sodium absorption was related negatively with Zn levels higher than 117.1 ppm in 2A/OR but positively in 2A/DR (Table 5; Figure 3). Such complexity in the relationship could be attributed to the variation in the interactions among major and micro nutrients caused by their relative abundance, variety and rootstock.

Author’s contributions

The corresponding author was involved in formulating the research programme, processing the data collected by other authors, graphical presentation of the results, interpretation of results and preparing the manuscript. All authors read and approved the final manuscript.

Acknowledgments

The authors are grateful to the Office Bearers and the Chairman, Central Research Committee of The Maharashtra Grape Growers’ Association for facilitating the conduct of the Survey; and the members of the research Advisory Committee for their suggestions and guidance in conducting the research. 

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