Mg application affected photosynthate production and distribution
Cherry tomato yield and dry matter accumulation were significantly affected by solution Mg concentration, which is consistent with the findings of Nzanza (2006) [26]. Increased yield and dry matter accumulation in response to proper Mg application was also observed by Hao and Papadopoulos (2003), who reported decreased fruit yield in the late growth stage at 0.82 mM solution Mg supply in rockwool blocks [4]. Moreover, in the later studies, Hao and Papadopoulos (2004) explained it by decrease of biomass and fruit biomass allocation in the low Mg treatment [15]. We also observed that lower plant DW and HI reduced yield in response to low Mg application (< 1 mM).
The photosynthetic rates of the cherry tomato plants decreased significantly in the 0 and 0.5 mM treatments. A previous study reported that the middle and bottom leaves of cherry tomato plants grown in a soilless production system showed leaf chlorosis under Mg starvation, losing about 50% of their photosynthetic capacity [4]. Impairment of sugar metabolism, photosynthetic CO2 fixation, and stomatal conductance were reported by Cakmak et al. (1994) and Fischer et al. (1998) in bean [27] and spinach [28] plants, and Andersson (2008) demonstrated that the involvement of rubisco in CO2 fixation was adversely affected by poor Mg supply [29].
The decrease of HI among cherry tomato plants observed under lower Mg supply in this study indicates the suppression of assimilate distribution to fruits. Sugar accumulation in source organs and the decline of its distribution to sink tissues have been reported previously. Hermans et al. (2004) found that sucrose accumulated in the most recently expanded sugar beet leaves before any loss of photosynthetic activity under Mg deficiency treatment [13]. Farhat et al. (2016) attributed it to preference of Mg transported to source leaves to prevent severe declines in photosynthetic activity [19]. Mg starvation seems to have a direct detrimental effect on function and/or structure of phloem loading [19, 27, 30, 31].
Relationships between leaf SPAD reading, photosynthetic rate, plant DW and leaf Mg concentration
Leaf Mg concentrations increased continuously as solution Mg levels increased in this study. A former study of Sulla carnosa plants also showed increased leaf Mg concentrations, to 2.5-, 7-, and 25- fold that of the control (0 mM Mg treatment) in 0.01, 0.05, and 1.50 mM Mg treatments, respectively [11]. In this study, the linear-with-plateau model illustrated the relationship between SPAD reading and leaf Mg concentration at the first and second harvests, and the critical leaf Mg concentrations for SPAD reading was about 4.67 and 5.52 g kg-1 in these periods. SPAD reading is an indicator of leaf chlorophyll concentration, which determines photosynthetic rate to a great extent [19]. So photosynthesis rates also fitted this model, and the critical leaf Mg concentration for photosynthesis rates was about 4.41 and 5.01 g kg-1 at the first and second harvests. A previous report indicated that maintenance of normal plant growth requires 4.0-6.0 g kg-1 leaf Mg concentration in tomato plants at anthesis, and the marginal concentration in first harvest period was 3.0 g kg-1 [32]. The linear-with-plateau model was applied to dry matter formation too, and the critical leaf Mg concentration was about 4.38 and 4.50 g kg-1, slightly lower than those for the photosynthesis rate. Similarly, dry matter accumulation in Pinus radiata was shown to be inhibited by Mg deficiency [33]. Hauer-Jákli and Tränkner (2019) confirmed 3.9 g kg-1 as the critical leaf Mg concentration for tomato dry matter accumulation [34] based on the results of Kasinath et al. (2014), which was lower than this study [35]. The different critical leaf Mg concentrations among SPAD reading, photosynthesis rate and plant dry matter accumulation indicated that sufficient Mg supply can guarantee the chlorophyll concentration and the production of photosynthates, which was consistent with the result that plant growth reduction appears as a later response compared with chlorophyll content decrease to Mg deficiency [14]. Clear relationships were observed between SPAD reading, photosynthesis rate, plant dry matter accumulation and leaf Mg concentration. It may be explained by the adequate Mg supply during initial growth stages [34]. These results clearly demonstrate the importance of Mg supply in maintaining strong photosynthesis to produce cherry tomato dry matter.
Two-side effects of Mg application on the plant K and Ca content
In the second harvest period, the plant K and Ca content of cherry tomato was first increased with Mg concentration in solution before 1 mM and then decreased. It was indicated by the plant dry matter accumulation and the leaf K and Ca concentration.
The plant dry matter accumulation increased first with solution Mg concentration increased but decreased when Mg treatment concentrations over 8 mM and 4 mM at the first and second harvest periods. The positive effects of Mg nutrient supply on plant growth have been discussed extensively [33, 34, 36], the inhibitive effects observed in this study have rarely been reported due to the difficulty of detecting toxicity symptoms, even at high concentrations [37]. The inhibitive effect of high Mg supply on plant dry matter accumulation was caused by slight decreases in the photosynthetic rate at first and second harvest. A similar effect was observed by Rao et al. (1987), who found that net photosynthesis was inhibited to a much greater extent in sunflower plants with a high Mg2+ content, particularly during dehydration [38]. Moreover, Shaul (2002) and Koch et al. (2019) associated this decrease with K+ transport inhibition from the cytosol to the stroma, disequilibration within the chloroplast and interference in transport events across the tonoplast [9, 18].
Low leaf K and Ca levels among high Mg supply treatments indicate antagonistic effects among these cations [24]. When solution Mg concentration was higher than 4 mM, the leaf K concentration was lower than 38.0 g kg-1, which might induce K deficiency [32, 39]. Leaf Ca concentration also decreased in higher Mg supply treatments, but was higher than 10 g kg-1 [32]. The response of fruit Ca concentration was more sensitive than that of K to Mg concentrations in this study, consistent with the results of Marschner (2012), who reported seven-fold higher K distribution than Ca distribution in pea seeds [24], and Karley and White (2009) also noticed this phenomenon [40].
K application influenced cherry tomato growth by regulating plant Mg and K
Antagonistic effects of K on Mg, especially under inadequate Mg supply conditions, are a crucial factor influencing Mg-related functions in several crops, including tomato [26], sugarbeet [41], green bean [42], potato [43], rice [44], grape [45] and apple [46]. The present study showed that an increasing K concentration in solution adversely affected the leaf Ca and Mg concentrations. Leaf Mg and Ca were lower than 4.7 and 10.0 mg kg-1 when solution K concentrations exceeding 17 mM under the 2 mM solution Mg supply, indicating Mg (from the former experiment in this study) and Ca deficiency [32]. However, K deficiency may have occurred when solution K concentration was less than 12 mM because leaf K concentration was lower than 38.0 mg kg-1 [32, 39]. These findings may explain the influence of K supply levels on total yields and plant DWs. Which was in line with Yurtseven et al. (2005), who reported that significant yield increases with increasing K application [47]. However, Nzanza (2006) found that none of the applied K treatments had any significant effect on marketable tomato yield [26]. The difference could be explained by the maximum K supply concentration, 9 mM in the study by Nzanza (2006) and 27 mM in this study [26].
Leaf K, Ca, and Mg concentrations are regulated by the K concentration in solution, as well as plant K, Ca, and Mg uptake. Ali et al. (1991) found that K, Ca, and Mg leaf contents in tomato decreased to 38%, 45%, and 67% of that of control plants under low K, low Ca, and low Mg supply, respectively, and that leaf, stem and petiole dry matter also decreased significantly [48]. Another study reported that rice shoot DW decreased by 12.9% at high K/Mg ratios in solution, whereas root DW increased by 12.1% as sugar partitioning and root morphological parameters changed [44]. Toumi et al. (2016) also reported that Mg uptake was inhibited by increase of K/Mg in the nutrient solution in Vitis vinifera, but no significant differences in leaf Ca concentration were detected among treatments [45].
Mg and K management in soilless vegetable production
Since the functions of Mg in the production, partitioning, and utilization of plant photoassimilates are irreplaceable, adequate Mg supply in the rhizosphere is essential for high-productivity soilless vegetable production systems. According to our results, 1-4 mM Mg in solution is needed to ensure leaf Mg concentrations exceeding 4.67 g kg-1 at the early harvest and 5.52 g kg-1 at late harvest. Which can satisfy the requirements for optimized SPAD, photosynthesis rate and plant dry matter accumulation combined with high fruit yield. Those leaf Mg concentrations are slightly higher than that reported in a previous study, which demonstrated that tomato dry matter accumulation responded best at 3.9 g kg-1 plant Mg concentration [34-35]. However, excessive Mg concentrations (> 8 mM) in solution should be avoided due to the risk of adverse effects on photosynthesis. Toxic effects that impair crop growth and development also showed by Guo et al. (2015) when Mg concentration in soil solution was higher than 8.5 mM [17].
Mg deficiency is a common problem in growth media fertilized only with N, P and K [7, 19]. Consequently, harmonious crop-specific nutrient management requires further attention. Overuse of K fertilizer not only wastes K resources but also disturbs Mg uptake and reduces yield [49-50]. Therefore, K concentrations in soilless culture system should be managed to supply sufficient leaf K to achieve high yield, while avoiding Mg uptake suppression due to excessive K. Consta´n-Aguilar et al. (2014) observed that cherry tomato fruit dry matter was higher when K concentrations ranged from 10 to 15 mM [51]. The current study indicates that 12 mM K in solution is optimal, based on our nutrient uptake and photosynthate production results. We also established relationships among leaf K or Mg concentration with cherry tomato dry matter in this study, which may be useful for understanding the mechanisms of yield formation in soilless vegetable production systems.