The approach described here enabled the frequent addition of only 6 to 12 µM ammonium, which reduced pH fluctuations and resulted in a steady-state solution concentration of ammonium. This approach also facilitated the growth of multiple crops of lettuce without replacing the nutrient solution.
Automated control to stabilize pH.
Our system is not the first to use ammonium salts to help control pH in recirculating hydroponics, but we are the first to show its long-term viability over repeated plantings in a zero-discharge nutrient solution. Bosman et al. (2024) recently maintained ammonium in a nutrient solution through automated control. They first calculated an ammonium-to-nitrate ratio to control pH. They then used an algorithm to determine the amount of N to add to the system to maintain EC and dosed proportions of the ammonium and nitrate salt solutions to maintain pH. The pH was controlled to within ± 0.5 pH units of their target (pH 6.1), but they did not use an acid or a base to avoid accumulating undesirable ions. Our approach does not require continuous EC monitoring or a control algorithm, avoids the accumulation of ions, and facilitates more stable pH control.
The trends we observed in the nutrient solution pH were largely as expected. We maintained the total N concentration instead of the ammonium-N concentration, which led to larger pH swings in the ADP and AS treatments that received more ammonium during each pH dose. Despite the AS treatment receiving the lowest amount of ammonium in trials three and four (Table 2), we observed the largest pH swings in this treatment. Although the ADP treatment received similar amounts of ammonium, the additional P provided may have helped buffer the rapid pH changes (van Rooyen and Nicol 2022b). Although the total ammonium received among treatments in plantings two through four was similar, the ammonium in the ADP and AS treatments was received less frequently and in larger doses than the AN treatment. This shows the value of a low, steady-state approach to ammonium management compared with larger, infrequent additions.
The ammonium-to-nitrate ratio.
The literature is replete with discussions on optimal ammonium-to-nitrate ratios for plant growth. The optimum ratio varies among species but most studies indicate that some ammonium improves growth (Chen et al. 2024). We achieved up to 13% of the N delivered as ammonium (≈1:8 ratio) and saw no effect on yield compared with 4% ammonium. Our objective was not to optimize the ratio, but to stabilize pH, reduce ion accumulation, and better simulate field conditions.
Many hydroponic solutions, such as Hoagland’s original solution number one (Hoagland and Arnon 1938), use only nitrate to simplify pH control. However, Hoagland realized the value of ammonium and, 12 years later, published Hoagland’s solution number two with ≈7% ammonium-N (Hoagland and Arnon 1950).
Effect of solution EC on PCE.
Managing nutrients by mass balance allows the elements with active uptake to be drawn down to low levels in the solution, which results in a low solution EC (Langenfeld et al. 2022). Solution EC is often maintained at a set point, but this often results in excessive concentrations of some nutrients in the recirculating solution.
The highest PCE occurred in the first planting in all treatments, decreased in the second planting, and increased again by the fourth planting. There was no trend in tissue nutrient concentrations over time, which indicates the PCE changes were not due to a nutritional stress. The EC was 0.4 mS·cm−1 at the beginning of planting four compared with 1 mS·cm−1 at the beginning of planting one, but because there was no significant difference between the PCE in planting one and four, the change in PCE cannot be linked to a decreasing EC over the course of the study. The second and third crop cycles had a lower PCE, but these crops were grown in the summer months in the greenhouse when higher air temperatures and a higher vapor pressure deficit may have contributed to the reduced PCE.
Relationship between ions in solution and EC.
The nutrient solution EC is determined by the differential contribution of nutrient ions based largely on their concentration and square of their charge (Griffin and Jurinak 1973). The ADP treatment had the lowest concentration of calcium, magnesium, and S in solution among all treatments, which likely contributed to the lowest EC. Although the P concentration was the highest, dihydrogen phosphate is monovalent and was potentially overshadowed by the higher divalent ion concentrations. The AS treatment had the highest S concentration, but not the highest EC.
The higher concentration of nitrate maintained throughout the study in the AN treatment may have led to its slightly higher EC. Although a nitrate concentration of 20 mg·L−1 may seem low, this is more representative of field conditions and does not represent N stress. For example, van Rooyen and Nicol (2021) found no difference in the growth rate of hydroponic kale when the N concentration in the nutrient solution was maintained at 154 mg·L−1 compared with 1.4 mg·L−1, which is less than half the lowest concentration of nitrate-N measured in solution during any point in our study.
The initial EC was not maintained over time as plants were allowed to take up nutrients as needed. Maintaining EC often leads to ion accumulation over time, which we sought to avoid. The decreasing EC indicated that plants were healthy and had active nutrient uptake. This would have been difficult to observe if the EC was maintained. A lower EC is also more representative of field nutrient conditions.
The benefits of low ion concentration in solution.
The EC was ≈0.4 mS·cm−1 in the later crop cycles, yet there were no significant decreases in nutrient concentration in leaf tissue. This shows the value of controlling nutrients by mass balance rather than by a setpoint EC. Nutrient accumulation in solution and luxury uptake have no effect on yield if adequate concentrations of nutrients are provided (Adler et al. 2000).
The ADP treatment resulted in higher P in solution and leaf tissue than the other treatments. The absence of a yield increase indicated that P was not limiting and did not benefit from luxury uptake. Penn et al. (2022) also found elevated P tissue concentrations in corn with increasing P concentrations but found no significant yield changes. Despite the elevated solution concentrations of S in the AS treatment, there was no difference in leaf tissue sulfur concentration. This indicates that lettuce does not take up S beyond that required for normal growth. At the high S concentrations measured in our study, the plants would have excluded S from uptake.
Because there was no difference in yield among treatments, excess nutrients were unnecessary and could lead to precipitation. The higher P concentration in the ADP treatment was correlated to a lower Fe concentration in both the nutrient solution and the young leaf tissue. This may be due to the precipitation of Fe as iron (III) phosphate. Amiri and Sattary (2004) studied nutrient solubility in solution culture and reported a 20% loss of Fe and a 15% loss of P from precipitation when their concentrations were doubled, which is similar to the decrease in Fe in solution that we observed in the final planting. It is thus beneficial to maintain low P concentrations to reduce Fe precipitation, which can lead to decreased bioavailability and iron chlorosis (Parry and Bugbee 2017).
Carbon partitioning to roots.
A higher percent root mass is often indicative of a lower concentration of nutrients in solution where a plant partitions more energy toward root growth to maximize nutrient uptake (Thornley 1972). The percent root mass in the final two plantings was slightly higher than reported in the literature. Sakamoto and Suzuki (2015) measured a percent dry root mass ranging from 13% to 15% for hydroponic lettuce ‘Red Wave’ and Li et al. (2018) found a percent root mass of ≈10% in hydroponic lettuce ‘Dasusheng’ and ‘Nenglv Naiyou’. In our study the first two plantings had a lower percent root mass. The higher nutrients in the first planting may have resulted in less energy partitioned to root growth, although the second crop cycle had a lower EC and a similar percent root mass. This indicates that changes in percent root mass may not be fully explained by concentrations in the nutrient solution.
Low pH is also a stress. The higher percent root mass in the last two crop cycles may have been caused by decreased pH. The replicates with the largest pH decreases were associated with the highest percent root masses.
Nitrification potential.
Nitrification is common in the field but is minimal in liquid hydroponics. Most nitrifying organisms need a solid substrate to grow, which is absent in liquid hydroponics. Padgett and Leonard (1993) measured a nitrate accumulation of 0.35 mM nitrate in sand culture with 2 mM ammonium, but nitrate accumulation was less than 1 µM (0.001 mM) in liquid culture. Muhlestein (2001) reported a nitrification rate of ≈250 µM·d−1 in a nutrient solution containing isolite, but when plants were added, they likely competed with the bacteria for ammonium uptake and nitrification was not detectable. Collectively these studies indicate that nitrification is minimal in liquid hydroponics without a substrate.