1 Introduction

Influence of soil physico-chemical characteristics on nutrient availability is well established (Truog, 1946; Russell, 1961). Absorption of a nutrient by a tree species depends not only on its availability, but also the relative contents of other nutrients (Emmert, 1959; Bergman et al., 1960), their ability to get adsorbed on to the root surface (Huffakar and Wallace, 1969; Wada and Weerasooriya, 1990), rootstock used(Smith and Wallace, 1956; Cook and Lider, 1964), affinity of the roots for nutrient ions (Asher and Ozanne, 1961; Downton, 1977) and nutrient interactions (Fageria, 2001; Wilkinson et al., 1999). Important chemical characteristics, which influence indirectly through their effects on nutrient availability and their interactions are, the organic carbon (Duxbury et al., 1989), pH (Sumner and Yamada, 2002), Electrical conductivity (Fisarakis et al., 2005), free calcium (Fageria, 2001) and exchangeable sodium percentage (Abrol et al., 1988). Rootstocks (Anna and Lajos, 2008; Antonio and Carlos, 2009; Marco et al., 2011) and the available nutrient status (Emmert, 1959; Bergman et al., 1960) were also shown to exert influence on the absorption of nutrients by grapevines. With this background, in a survey conducted in Thompson Seedless vines on their own roots and grafted on Dog Ridge and 110R rootstocks to study the variation, if any, in the absorption of nutrients under the influence of the soil chemical characteristics on different rootstocks, the ratio of petiole nutrient content to the available soil nutrient content was considered as the ‘nutrient absorption index’. Thompson Seedless is grown extensively in saline-alkali soils on these rootstocks and own roots in the tropical region of India. Results of the survey would guide in the nutrient management in soils with varying chemical characteristics on different rootstocks.

2 Materials and Methods

Thompson Seedless vineyards, 18 each on own root and Dog Ridge and 110R rootstocks, in the age group of 5-7 years were identified for this study. Vines were raised on vertisols with varying levels of available nutrients and receiving varying levels of nutrients. The organic carbon content of the soils in the selected vineyards was 2.57 ± 1.08 per cent, pH 7.76 ± 0.48, EC 0.61 ± 0.42 dSm-1 , CaCO3 16.2 ± 4.2 per cent and Exchangeable sodium percent 7.77 ± 1.82. All the vines selected for the study were planted at 3.0 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. N, P, K, Ca, Mg, S, Na, Fe, Mn, Zn, Cu contents in petiole and soil samples; and organic carbon, pH, EC, CaCO3 and ESP in soil samples were estimated following the standard analytical methods suggested by the AOAC. The ratio of petiole nutrient content to that of soil was calculated in order to normalize the nutrient absorption and termed “nutrient absorption index” for each nutrient.

Correlations of soil chemical characteristics with the absorption index of nutrients, multiple regression of nutrient absorption index on chemical characteristics and the quadratic equations for the significant relation of soil chemical characteristics with nutrient absorption index were calculated by the Microsoft Excel data analysis package. Significance of the coefficients of the soil chemical characteristics in each regression equation was tested by their respective t-stat values. Optimum value of each chemical characteristic (X-opt.) was calculated by the formula; X-opt= –b/2c, where ‘b’ is the coefficient of X and ‘c’ the coefficient of X2 in the respective quadratic function and the maximum absorption index for each nutrient (Y-max.) was derived by substituting the X values in the corresponding function.

3 Results and Discussion

Multiple regression analysis revealed that, all the soil chemical characteristics, namely organic carbon (OC), pH, EC, CaCO3 and ESP together influenced the absorption of all the nutrients studied, except Ca and Cu in vines on own root; N, P and Cu on Dog Ridge; and Ca, Mg, Na and Cu on 110R. Absorption of K, S, Fe, Mn and Zn was influenced on all roots, whereas Mg and Na on own root and Dog Ridge; N and P on own root and 110R; and Ca on only dog Ridge. Nutrient absorption was influenced most on own root followed by Dog Ridge and 110R roots as evidenced by their determination coefficients.  N followed by Zn, P and K were the most influenced nutrients on own root, while K, Mn and Zn on Dog Ridge; and Fe, N, K and Zn on 110R. All the soil chemical characteristics were ineffective in the absorption of Cu in vines on any root. Roots of Thompson Seedless and 110R were similar in their absorption of Ca but differed in Mg absorption. While Mg absorption was not influenced on 110R, it was on own root. Dog Ridge roots differed with other roots in respect of the absorption of N and P; while 110R in Na (Table 1). Lack of influence of the soil chemical characteristics in the absorption of any nutrient on any root could be due to the greater availability of the nutrient and/or the higher affinity of the roots for it. Organic carbon contributed positively towards the determination of absorption of N, P, K and Fe on own root and K on Dog Ridge, but negatively in case of Mn and Zn on Dog Ridge; and N and K on 110R. pH contributed positively in the determination of Ca, Mg, Na and Zn on own root, Ca and Na on Dog Ridge; and N and K on 110R, while negatively in case of P, K, S, Fe and Mn on own root; K and Fe on Dog Ridge; and Fe and Mn on 110R. EC contributed positively in the determination of absorption of S on own root; Mg, Fe, Mn and Zn on Dog Ridge; and Fe, Mn and Zn on 110R, but negatively in case of Ca and Mg on own root; Na on Dog Ridge and P on 110R. ESP contributed positively in the case of Mn on own root and P on 110R, but negatively in case of N on own root and Zn on 110R (Table 1). Variation in the contribution of chemical characters on different roots is elucidated by the variation in the absorption of nutrients under their influence in the following paragraphs.

3.1 Effect of organic carbon

When the individual effect of soil chemical characteristic on the absorption of nutrients on different roots is considered, organic carbon (OC) content of soil influenced the absorption of total nitrogen and sulphur in vines on their own root. None of the nutrients was influenced by it either on Dog Ridge or 110R rootstocks (Table 2). While the absorption of total nitrogen increased steadily with increasing levels of OC on own root, it decreased on 110R and increased on Dog Ridge up to 3.03 per cent (Table 3). Response of Dog Ridge in the absorption on N to additional levels of OC was more than own root up to 3 per cent. 110R was more efficient than Dog Ridge under low levels of OC up to 1.8 per cent and own root up to 2.7 per cent, whereas Dog Ridge was more efficient above1.8 per cent (Figure 1). OC being the indicator of organic matter in the soil (Jackson, 1958) and promoter of N absorption (Greenland and Nye, 1959), the results imply that its requirement is less on 110R rootstock for N absorption.

The significant negative relationship of sulphur absorption with soil OC on own root but non-significant one on other roots (Table 2) indicate that, sulphur absorption by Thompson Seedless root was more independent of soil OC than other roots. Increasing levels of OC up to 2.06 per cent were associated with reduced sulphur absorption on own root and steady reduction on 110R, but increased absorption up to 3.3 per cent on Dog Ridge (Table 3). Thompson Seedless root were more efficient than the other roots in S absorption at all levels of OC, while 110R was more efficient than Dog Ridge up to 2.0 per cent (Figure 1). Importance of OC in the absorption of S lies in the fact that 60 per cent of the sulphur taken up by the plants being derived from carbon-bonded sulphur fractions. Positive response of Dog Ridge roots indicates their dependence on organic carbon for S absorption.

3.2 Effect of soil pH

Soil pH influenced negatively the absorption of P and S on own root; and Ca and S on Dog Ridge but none of the nutrients on 110R (Table 2). Phosphorus absorption reduced with increasing levels of pH on own and Dog Ridge roots respectively up to 8.13 and 8.19 (Table 2) and increased thereafter. Own root was more sensitive to soil pH in the absorption of P compared to other roots. Though P absorption increased with pH up to 7.58 on 110R, the response was poor. Efficiency of P absorption was highest on own root (Figure 2). The pattern of P absorption in relation to pH on own root and Dog Ridge was in accordance with the P availability that was maximum at near neutrality (soil pH 6.5–7.5), and decreased at higher and lower pH (Truog, 1946; Sumner and Farina, 1986).

Soil pH also influenced calcium absorption significantly on Dog Ridge, but not on other roots (Table 2). Calcium absorption was reduced with increasing levels of pH up to 7.51 on Dog Ridge, 7.88 on own root and 7.23 on 110R (Table 3). Own root were more sensitive to soil pH than other roots. Rootstocks were more efficient in the absorption of Ca at neutral pH. Higher pH levels (above 8) were favourable for Ca absorption (Figure 2). This could be attributed to alkalinity (pH 7.76 ± 0.48) coupled with high free calcium content (16.2 ± 4.2) of the vineyard soils. Variation in the response and sensitivity of different roots in the absorption of Ca under the influence of soil pH could be due to root preference and/or the uptake other cations.

Sulphur absorption reduced significantly with increasing levels of soil pH on Dog Ridge (Table 2). While it increased up to a pH of 7.79 and reduced thereafter on own root, reduced up to 8.48 and increased marginally thereafter on Dog Ridge (Table 3). Dog Ridge root were more efficient than the rest two and own root was more sensitive to changes in soil pH, while 110R root was independent of pH in S absorption (Figure 2). Within the pH range of vineyard soils in the present survey, absorption of Ca and S were opposite to each other on all types of root (Figure 2). This indicates the mutual antagonism between Ca and S under the condition of pH and free calcium contents of the vineyard soils.

3.3 Effect of soil electrical conductivity

Electrical conductivity of the soil (ECe), among the chemical characteristics, influenced the highest number of nutrients studied. It influenced positively the absorption of N on own root; S, Mn and Zn on Dog Ridge and Fe on 110R (Table 2). N absorption increased steadily with increasing levels of ECe on own root, while increased up to 0.62 and decreased thereafter on Dog Ridge but reduced up to 1.12 and increased marginally thereafter (Table 3). Own root was more responsive to changes in ECe in N absorption. While 110R was more efficient in N absorption at ECe level less than 0.5 dSm-1 and Dog Ridge at 1.0 dSm-1, own root were more efficient beyond 1.0 dSm-1 (Figure 3).

Absorption of S increased up to ECe of 0.88dSm-1 and reduced thereafter on own root, but increased steadily but marginally with increasing levels of ECe on the rootstocks (Table 3). Own root were more efficient in S absorption in the ECe range of 0.5 -1.5 dSm-1, while the rootstocks were more efficient beyond 1.5dSm-1 (Figure 3).

Absorption of Fe reduced with increasing levels ECe up to 0.81dSm-1 and increased thereafter on own root (Table 3). Though the response was positive and steady on 110R, the rate of increase in Fe absorption to unit increase in ECe was higher on own root than on 110R beyond 1.0dSm-1, consequently own root was more efficient than the other roots beyond 1.5dSm-1 (Figure 3).

Manganese absorption increased steadily with increasing levels of ECe up to1.76dSm-1 on Dog Ridge, but decreased up to 0.88dSm-1 and increased thereafter on own root (Table 3). Own root were more efficient in Mn absorption at lower levels of ECe up to 0.5dSm-1, whereas Dog Ridge was more efficient at higher levels beyond 0.5dSm-1compared to other roots (Figure 3).

Absorption of Zn decreased marginally up to 0.59dSm-1 of ECe and increased sharply thereafter, whereas it increased at a lower rate after 0. 59 dSm-1 on own root (Table 3). Own root were more efficient up to 1.1dSm-1, while Dog Ridge beyond that, when compared to other roots (Figure 3). The positive relationship of Mn with ECe and negative relationship of Zn on Dog Ridge root indicated their mutual antagonism on this rootstock. Similarly, the nature of response curves of nutrients to ECe was the pointer for mutual antagonism between S and Mn and synergism between Fe and Mn on own root. Variation in the nutrient interactions on different roots was attributed to root affinity for nutrients (Downton, 1977; Asher and Ozanne, 1961). Relative contents of soluble salts of chlorides, sulphates and bicarbonates of potassium, calcium, magnesium and sodium in the soil contribute to the level of ECe. The specific ion concentration contributing to ECe, its interaction with other nutrient ions seem to be the reasons for variation in the absorption of nutrients by different roots under varying levels of ECe.

3.4 Effect of free calcium content of soil

Free calcium CaCO3 content of soil influenced K absorption negatively on own root and Dog Ridge, but positively that of Ca on Dog Ridge. None of the nutrient absorption was influenced on 110R (Table 2). Negative influence of CaCO3 was observed from 7.38 per cent onwards on own root and from 8.39 per cent on Dog Ridge (Table 3). Own root was more efficient in K absorption than Dog Ridge at all the levels of CaCO3 in the present study. 110R root were more efficient than the rest at higher levels (>17 per cent) – Figure 4. Antagonistic effect of Ca on K is a well established fact (Wilkinson et al., 1999). Lack of antagonism on 110R could be due to strong affinity of its root for K (Kalbhor et al., 2017).

Calcium absorption decreased marginally with increasing levels of CaCO3 up to 6.75 per cent on dog Ridge and 12.5 per cent on 100R. It increased steadily thereafter on Dog Ridge but marginally on 110R. Ca absorption increased sharply from 8.0 to 17.4 per cent CaCO3 (Table 3). Own root were more sensitive to CaCO3 levels in Ca absorption with a threshold level of 8.0 per cent.  Rootstocks were more efficient than own root at CaCO3 levels < 10 per cent, while the later were more efficient between 10-20 per cent. Dog Ridge continued to be more efficient even beyond 20 per cent (Figure 4). Non significant relationship of Ca absorption with CaCO3 on own root and 110 R indicates that, these roots have more affinity for those cations which are antagonistic to Ca.

Absorption of sodium decreased increased with increasing levels of CaCO3 up to 14.9 per cent and increased thereafter. It increased up to 20.4 per cent on own root and 14.4 on 110R. Dog Ridge roots tended to absorb more Na up to 10 per cent CaCO3, while own root beyond that level. They were more efficient in restricting the absorption of Na in the CaCO3 range of 10-20 per cent, while 110R roots at all levels. Both rootstocks were at par in the absorption of Na in the range of 12-17 per cent (Figure 4). Thus the ability of restricting the absorption of Na by rootstocks was dependent on the free calcium levels in soil.

3.5 Effect of exchangeable sodium percent in soil

ESP influenced the nutrient absorption on own root only. It influenced the absorption of N, P and K negatively but Fe positively (Table 2). N absorption decreased steadily with increasing levels of ESP reaching null at 14.5 per cent on own root and 15.5 on 110R, but increased up to 8 per cent, reducing thereafter, reaching null at 15 per cent on Dog Ridge (Table 3). Own root were more efficient than the rootstocks up to 6.0 per cent, while Dog Ridge beyond that (Figure 5). Reduction of N use efficiency could be due to denitrification and volatile losses caused by poor drainage resulting from high levels of sodium in medium clay soils (Abrol et al., 1988).

Absorption of P decreased steadily with increasing levels of ESP up to 14.1 per cent on own root, but increased up to 8.23 and 8.15 per cent respectively on Dog Ridge and 110R and reducing thereafter (Table 3). Own root were more efficient in P absorption up to 7.0 per cent ESP. Dog Ridge was slightly more efficient than the rest between 10 1nd 14 per cent (Figure 5). Low P absorption was attributed to higher pH levels, since positive correlation between PH and ESP was observed (Gupta et al., 1981).

Potassium absorption increased up to 4.89 and 7.76 per cent  ESP respectively on own root and 110R and reduced thereafter; but reduced up to 10.4 per cent on Dog Ridge increasing thereafter (Table 3). Own root were more efficient than the rootstocks between 3.5 – 7.76 per cent ESP, while Dog Ridge beyond 11.5 (Figure 5). Reduction in K absorption could be attributed to the antagonistic effect of Na on K (Fisarakis et al., 2005; Shikhamany and Sharma, 2008).

Absorption of Fe reduced with increasing levels of ESP up to 7.27 per cent and increased beyond it (Table 3). Own root were less efficient than others between 6 and 10 per cent ESP, during which range 110R was more efficient (Figure 5). Positive effect of ESP on Fe absorption could be due to suppression of the absorption of Ca which is antagonistic to Fe.

Variation in the efficiency in nutrient absorption with the rootstock at different levels of soil chemical characteristics could be attributed to the effects of specific ion contributing to the chemical character, its interaction with other nutrient ions, their relative abundance and the preferential absorption by the roots, soil pH and soil physical properties. The very fact that crops/ genotypes are identified in relation to soil pH, salinity and exchangeable sodium have been identified (US salinity laboratory Staff, 1954), itself indicates the variation in the response of genotypes due to many reasons including nutrient absorption.

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