Abstract
In soilless greenhouse tomato cultivation, daily transpiration and irrigation
demand are largely governed by solar radiation, while irrigation-solution electrical
conductivity (EC) used for salinity management may further modulate plant water
use. This study developed a low-input, radiation-driven modeling approach to predict
daily irrigation demand under contrasting water–salt management scenarios. Two
tomato cultivars were grown under four treatments: conventional baselines (CK1,
CK2) and regulated scenarios combining irrigation volume with solution EC
(low-water high-EC, TK; high-water moderate-EC, TC). Daily irrigation volume (I)
and drainage were recorded, and daily cumulative radiation (G) was derived from
photosynthetically active radiation (PAR). Within each treatment, we compared a
radiation-only baseline model with an EC-adjusted model and evaluated predictive
performance using 5-fold blocked time-series cross-validation. Results showed strong
positive correlations between G and I across all treatments (p < 0.001). The
EC-adjusted models achieved cross-validated root-mean-square errors (RMSE) of
0.815–1.393 L d⁻¹ per trough and Nash–Sutcliffe efficiencies (NSE) of 0.407–0.730.
Incorporating EC yielded a small but consistent improvement under the TK scenario
(ΔRMSE = −0.014 L d⁻¹; ΔNSE = +0.019), whereas its effect was negligible or
slightly negative under CK1, CK2, and TC, highlighting scenario dependence. Our
radiation-driven framework, with an optional EC correction, offers a practical and
scalable tool for daily irrigation forecasting and supports integrated water–salt
management in soilless greenhouse tomato production.
Keywords
greenhouse tomato; irrigation demand; solar radiation; electrical
conductivity; substrate cultivation; irrigation scheduling
1. Introduction
1.1. Greenhouse tomato production and irrigation challenges
Tomato (Solanum lycopersicum L.) is a globally significant protected
horticultural crop, whose economic viability hinges on yield stability, fruit quality,
and resource-use efficiency [1,2]. With the expansion of greenhouse cultivation,
management strategies have evolved from a sole focus on yield toward integrated
approaches that harmonize productivity, quality enhancement, and water conservation
[3,4]. Nevertheless, greenhouse tomato systems remain heavily reliant on irrigation
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
and fertigation, rendering water management a critical—and often limiting—factor in
the context of escalating water scarcity and rising production costs [5–7].
In practice, irrigation scheduling in greenhouses frequently depends on grower
experience or fixed timetables, which seldom account for rapid daily fluctuations in
crop water demand. Suboptimal irrigation not only curtails water-use efficiency but
also perturbs root-zone salinity and nutrient dynamics [8,9], thereby influencing
photosynthetic performance, yield formation, and key fruit quality attributes such as
soluble solids and nutritional composition [10–12]. Hence, developing irrigation
strategies that dynamically respond to environmental variability while safeguarding
yield and quality remains a pivotal challenge in modern greenhouse tomato
production.
1.2. Radiation-driven irrigation demand in greenhouse systems
Among environmental drivers, solar radiation exerts a dominant influence on
canopy transpiration and carbon assimilation in greenhouse tomatoes [13,14]. Daily
variations in photosynthetically active radiation (PAR) directly modulate stomatal
conductance and transpiration rates, establishing radiation as a primary determinant of
short-term irrigation demand [15–17]. Unlike temperature and humidity, which can be
partially regulated in controlled environments, incident radiation is largely governed
by external weather conditions, leading to pronounced day-to-day variability in crop
water requirements [18,19].
Empirical relationships between daily cumulative radiation and irrigation water
consumption have been consistently demonstrated in greenhouse tomatoes, furnishing
a practical foundation for radiation-based estimation and scheduling approaches
[13,20,21]. Compared with mechanistic evapotranspiration models or data-intensive
machine-learning techniques, empirical radiation-driven models offer distinct
advantages in simplicity, robustness, and operational feasibility—attributes especially
valuable for daily irrigation scheduling in commercial settings [22–25]. However,
most existing radiation-based studies have been conducted under uniform fertigation
regimes, seldom explicitly addressing potential interactions with root-zone salinity
management [26].
1.3. Electrical conductivity and water–salt interactions
In soilless and substrate-based greenhouse tomato systems, the electrical
conductivity (EC) of the nutrient solution is a key operational variable that shapes the
root-zone water–salt environment [27–29]. Adjustments in EC alter osmotic potential
and ion concentrations around roots, thereby influencing water uptake, transpiration,
and assimilate partitioning [30,31]. Numerous studies indicate that moderate elevation
of EC, or controlled deficit irrigation, can enhance fruit quality traits such as soluble
solids and sugar-acid ratio, though excessive salinity may suppress yield and overall
plant water consumption [32–34].
Critically, the effects of EC on crop water use are not independent of irrigation
supply level. Under differing water-availability backgrounds, identical EC conditions
may exert contrasting influences on transpiration and irrigation demand, owing to
shifts in plant water status and root-zone hydraulic properties [35,36]. Recent work
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
emphasizes the importance of integrated water–salt regulation rather than treating EC
as a static background parameter [37–40]. Despite this, most irrigation-scheduling
frameworks continue to neglect EC-dependent adjustments in daily irrigation demand,
particularly in soilless cultivation systems.
1.4. Research gap and objectives
Although radiation-driven approaches are widely employed to estimate
greenhouse irrigation demand, prevailing frameworks rarely incorporate EC-related
adjustments under contrasting water–salt management scenarios. This omission
constrains their applicability for integrated irrigation strategies that simultaneously
aim for water conservation and fruit quality improvement.
Therefore, this study aimed to develop a simple, practical radiation-based
Method
for daily irrigation demand estimation under varying electrical conductivity
scenarios in substrate-grown greenhouse tomato. Specific objectives were to:(i)
quantify the relationships between daily cumulative radiation and irrigation demand
under different EC management backgrounds;(ii) elucidate how EC modifies
radiation-driven water-demand responses; and (iii) evaluate the implications of
EC-adjusted irrigation scheduling for yield stability and fruit quality.
By emphasizing operational feasibility and employing treatment-wise blocked
time-series cross-validation, this work provides a management-oriented irrigation
framework that complements more complex modeling approaches reported in the
literature.
2. Materials and Methods
2.1. Experimental Site and Environmental Conditions
The experiment was conducted from November 2023 to May 2024 in Glass
Greenhouse A2 at the Facility Agriculture Research Base of the Academy of
Agricultural Planning and Engineering, Ministry of Agriculture and Rural Affairs,
located in Yongqing County, Langfang City, Hebei Province, China. The greenhouse
covered approximately 5000 m² and was equipped with automated
environmental-control systems (internal/external shading, fan-pad cooling, heating,
supplemental lighting, and ventilation). The daytime temperatures were controlled
between 22°C and 26°C, and night-time temperatures ranged from 18°C to 22°C.
Relative humidity was maintained at 60%-70%, and environmental data (such as
radiation and temperature) were continuously recorded using an on-site automatic
weather station.The complete daily environmental monitoring and irrigation records
are provided in S1 Dataset.
2.2. Plant Materials and Cultivation Method
Test varieties were ‘Jingdan No. 8’ (cherry tomato) and ‘Jingcai No. 8’
(strawberry tomato). Soilless substrate cultivation used a 3:1 (v/v) coconut coir:perlite
mix. Each substrate trough measured 1.0 m × 0.15 m × 0.12 m (length × width ×
height), with four plants per trough (plant spacing 30 cm, row spacing 55 cm) (Figure
1). Drip irrigation was applied using municipal tap water, with nutrient solution
adjusted for pH and EC before irrigation. Prior to planting, coconut coir was fully
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
hydrated and pretreated with standard nutrient solution to minimize initial water-salt
variability.
Figure 1. Schematic diagram of the substrate cultivation trough and drip irrigation setup for
greenhouse tomato experiments.
2.3. Experimental Design and Treatments
A randomized block design was used with four water-fertilizer treatments:
conventional water-salt baseline for each variety (CK1 and CK2) and corresponding
regulated scenarios (low-water high-EC: TK; high-water moderate-EC: TC). Each
treatment had three replicates (12 plots total).
For model development, daily measurements were averaged across the three
replicate troughs within each treatment, yielding one daily observation per treatment
per day (approximately 180 days per treatment).
Considering varietal differences in water demand and nutrient management, a
“variety–water-salt scenario pairing” framework was adopted. ‘Jingdan No. 8’
received conventional baseline (CK1) and low-water high-EC (TK) to analyze supply
level and EC interactions under the same variety. ‘Jingcai No. 8’ received
conventional baseline (CK2) and high-water moderate-EC (TC) to examine EC
adjustments under enhanced supply. CK1 and CK2 served as practical baselines for
their respective varieties. Although initial design intended pH adjustment for CK2,
actual irrigation pH remained similar across CK1 and CK2 (Table 1); thus, CK2 was
positioned as the baseline for ‘Jingcai No. 8’ without treating pH as an independent
factor.
Table 1. Experimental treatment setup.
Treatment Variety Irrigation Volume
(L d⁻¹ per trough) EC (dS m⁻¹) pH
CK1 Jingdan No. 8 4.88 3.00 6.05
TK Jingdan No. 8 4.14 3.31 6.08
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Treatment Variety Irrigation Volume
(L d⁻¹ per trough) EC (dS m⁻¹) pH
CK2 Jingcai No. 8 4.53 3.21 6.08
TC Jingcai No. 8 5.65 3.18 6.28
Target irrigation volumes, EC, and pH were maintained via flow meters (continuous recording)
and online/manual adjustments for stability throughout the trial.
2.4. Measurements and Data Collection
Irrigation volume was recorded in real time by flow meters; drainage was
collected and summarized daily. Nutrient solution EC and pH were measured using
conductivity (INESA DDS-307A) and pH meters (PHS-3C), calibrated periodically.
Light conditions were monitored with quantum sensors (LI-190R, LI-COR,
USA) placed at greenhouse top, recording PAR at 1-min intervals. Daily cumulative
radiation (G, MJ m⁻² d⁻¹) was integrated from PAR. Specifically, PAR (μmol m⁻² s⁻¹)
was integrated to daily light integral (DLI, mol m⁻² d⁻¹) and converted to energy units
using a conversion factor of 0.218 MJ per mol photons . At the daily aggregation
scale used for modeling, no missing values were present; therefore, no interpolation or
exclusion was required. As a prespecified quality-control rule, if PAR gaps had
occurred, gaps ≤10 min would have been linearly interpolated, whereas days
with >5% missing PAR would have been excluded from modeling.
Photosynthetic parameters (net photosynthetic rate Pn, transpiration rate Tr,
stomatal conductance Gs, intercellular CO₂ concentration Ci) were measured using a
portable system (LI-6400XT) on three representative plants per treatment (three
measurements per plant, averaged). Photosynthesis measurements are available in S4
Dataset.
Yield was recorded at the plot level (one cultivation trough per replicate) at each
harvest (fruit number and total fresh weight). Yield was expressed as g plot⁻¹ per
harvest (and kg plot⁻¹ for seasonal cumulative yield). Per-harvest yield and fruit
number data are available in S2 Dataset.
2.5. Data Processing and Statistical Analysis
Data were organized in Microsoft Excel and analyzed using SPSS (version 25.0).
Treatment effects were assessed via one-way ANOVA at α = 0.05, with Duncan’s
multiple range test applied for post-hoc comparisons. Pearson correlation coefficients
were computed to evaluate linear associations, with significance levels denoted as *p
< 0.05, **p < 0.01, and ***p < 0.001.
To quantify the combined influence of radiation and irrigation-solution EC on
daily irrigation volume, an EC-adjusted linear regression model was fitted separately
for each treatment:
I = β0 + β1G + β2ECirr + ε (1)
A radiation-only baseline model was also fitted for comparison:
I = α0 + 𝛼1G + ε (2)
where I is daily irrigation volume (L d⁻ ¹ per trough), G is daily cumulative
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
radiation (MJ m ⁻² d⁻¹) integrated from PAR, ECirr is the electrical conductivity of
the irrigation solution (dS m⁻¹), βi and 𝛼i are regression coefficients, and ε is the
random error term.
Prior to final model fitting, a 3σ residual rule was applied within each treatment
to exclude outliers: an initial model was fitted, residuals were calculated, and
observations with |residual| > 3σ were removed. The model was then refitted on the
filtered dataset.
To evaluate out-of-sample predictive performance while accounting for temporal
autocorrelation, a blocked 5-fold time-series cross-validation was implemented. Daily
observations were ordered chronologically and partitioned into five contiguous time
blocks. In each fold, one block served as the validation set, while the remaining four
blocks constituted the training set. The 3σ outlier removal was applied solely to the
training data within each fold. Predictions from all folds were pooled, and
performance metrics—root-mean-square error (RMSE), mean absolute error (MAE),
and Nash–Sutcliffe efficiency (NSE)—were computed from the pooled out-of-sample
predictions.
3. Results
3.1. Correlations among Key Irrigation–Drainage Variables
Daily-scale relationships among irrigation, drainage, and radiation variables were
examined using Pearson correlations, including irrigation volume (I),
irrigation-solution electrical conductivity (EC irr) and pH, drainage volume (D),
drainage EC (EC dra) and pH, and daily cumulative radiation (G). The correlation
structure is illustrated in Figure 2 (underlying data in S1 Dataset).
Figure 2. Correlation matrix of key irrigation, drainage, and radiation variables at the daily scale.
Color intensity indicates correlation strength and direction. Asterisks denote significance (* p <
0.05, ** p < 0.01, *** p < 0.001). I, irrigation volume (L d⁻¹ per trough); D, drainage volume (L
d⁻¹ per trough); G, daily cumulative radiation (MJ m ⁻² d⁻¹); EC irr, electrical conductivity of
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
irrigation solution (dS m ⁻¹); pH irr, pH of irrigation solution; EC dra, electrical conductivity of
drainage solution (dS m⁻¹); pHdra, pH of drainage solution.
I was strongly and positively correlated with D (r = 0.70, p < 0.001), consistent
with the coupled dynamics of substrate water input and outflow. G was also positively
correlated with I (r = 0.65, p < 0.001), supporting the dominant role of
radiation-driven transpiration in determining day-to-day irrigation demand.
Among solution properties, EC irr was moderately negatively correlated with
irrigation pH (r = −0.32, p < 0.001). In addition, G was negatively correlated with
ECirr (r = −0.37, p < 0.001), suggesting that EC tended to be reduced on
high-radiation days, potentially as an operational adjustment to mitigate salt
accumulation risk.
3.2. Daily Irrigation Demand Prediction Models
Daily-scale empirical regression models were developed separately for CK1,
CK2, TK, and TC, using daily accumulated radiation (G) and irrigation water
electrical conductivity (EC irr) as predictors and daily irrigation amount (I) as the
response variable.
Drainage electrical conductivity (EC dra) was not statistically significant during
candidate-variable screening (p > 0.05) and was therefore excluded from the final
models.
CK1:
I = 2384.62 + 2065.40G - 421.35ECirr
R² = 0.79, n = 179; G showed a highly significant positive effect (p < 0.001),
while ECirr showed a significant negative effect (p = 0.025).
CK2:
I = 605.17 + 1687.12G + 259.51ECirr
R² = 0.71, n = 180; G showed a highly significant positive effect (p < 0.001).
The effect of ECirr was positive but marginal (p = 0.052).
TK:
I = 1252.60 + 995.08G + 360.98ECirr
R² = 0.52, n = 180; G showed a highly significant positive effect (p < 0.001),
and ECirr showed a significant positive effect (p = 0.032).
TC:
I = 3511.60 + 1555.54G - 191.25ECirr
R² = 0.61, n = 182; G showed a highly significant positive effect (p < 0.001).
The effect of ECirr was negative but not significant (p = 0.279).
Overall, the four models consistently indicate that G is the dominant driver of
day-to-day irrigation variation, whereas the direction and significance of ECirr
depend on the water management scenario.
3.3. Model Performance Evaluation
Model performance was evaluated using the coefficient of determination (R²),
root mean square error (RMSE), mean absolute error (MAE), and Nash–Sutcliffe
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
efficiency (NSE) (Table 2). RMSE and MAE quantify daily scheduling errors in
physical units (L d⁻¹ per trough), whereas NSE summarizes how well the model
reproduces day-to-day variability relative to the observed mean.
Table 2. Performance indicators of radiation-driven daily irrigation models under different
treatments.
Treatment n R² RMSE (L d⁻¹ per
trough)
MAE (L d⁻¹ per
trough) NSE
CK1 179 0.79 1.01 0.79 0.79
CK2 180 0.71 0.93 0.72 0.71
TK 180 0.52 0.75 0.57 0.52
TC 182 0.61 1.13 0.85 0.61
Across treatments, the fitted radiation-driven models achieved RMSE values of
0.75–1.13 L d⁻¹ per trough and MAE values of 0.57–0.85 L d⁻ ¹ per trough (Table 2),
indicating that daily irrigation demand was typically estimated within approximately
0.6–0.9 L d⁻¹. For the in-sample fitted models, NSE values numerically matched R²
because both metrics were computed as 1 − SSE/SST on the same dataset with an
intercept term; therefore, RMSE and MAE are particularly informative for operational
scheduling accuracy.
Performance differed among water–salt scenarios. CK1 exhibited the strongest
fit (R²/NSE = 0.79), suggesting a stable radiation–irrigation response under
conventional management for ‘Jingdan No. 8’. CK2 and TC showed intermediate
explanatory power (R²/NSE = 0.71 and 0.61, respectively), whereas TK had the
lowest explained variance (R²/NSE = 0.52), implying larger residual variability under
the low-water high-EC regime. Notably, TK also showed the smallest absolute errors
(RMSE = 0.75 L d⁻¹; MAE = 0.57 L d⁻¹), consistent with the lower irrigation supply
level limiting the magnitude of day-to-day fluctuations (Table 1).
In relative terms, RMSE corresponded to approximately 18–21% of the mean
daily irrigation volume across treatments, supporting the practical interpretability of
the error magnitudes for daily scheduling. Because in-sample metrics may overstate
predictive performance when temporal dependence is present, an out-of-sample
assessment based on blocked time-series validation is reported in the following
section.
3.4 Baseline Comparison and Blocked Validation
Within each treatment, a radiation-only baseline model was compared with an
EC-adjusted model using blocked 5-fold time-series cross-validation. Daily
observations were ordered chronologically and split into five contiguous time blocks;
in each fold, one block was held out for validation and the remaining blocks were
used for training. Validation predictions were pooled across folds, and RMSE, MAE,
and NSE were computed from the pooled out-of-sample predictions (Table 3).
Table 3. Blocked 5-fold time-series cross-validation performance of the radiation-only
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
baseline model and the EC-adjusted model (pooled across folds).
Note: RMSE and MAE are expressed in L d⁻¹ per trough; NSE is dimensionless. Δ indicates
EC-adjusted minus baseline; negative ΔRMSE/ΔMAE and positive ΔNSE indicate improvement.
The cross-validated results indicate that the incremental value of adding EC was
scenario dependent. Under the low-water high-EC scenario (TK), incorporating EC
reduced RMSE from 0.829 to 0.815 L d⁻¹ per trough and increased NSE from 0.422
to 0.441 (ΔRMSE = −0.014; ΔNSE = +0.019), reflecting a small but consistent
improvement. Although the magnitude of improvement was modest, the direction of
change (ΔRMSE 0) suggests that EC carries incremental predictive
information only under specific water–salt backgrounds. In contrast, EC adjustment
slightly degraded performance under CK1 and CK2 (ΔNSE = −0.014 and −0.008,
respectively) and produced a larger decline under TC (ΔRMSE = +0.080; ΔNSE =
−0.066). These comparisons support treating EC as a scenario-specific adjustment
factor—most useful under low-water high-EC management—rather than a universal
predictor that improves accuracy across all treatments.
Compared with in-sample fitting, NSE decreased under cross-validation, as
expected when predicting unseen time blocks, with baseline NSE ranging from 0.422
to 0.744 across treatments. Pooled observed–predicted relationships corroborate the
numerical comparison: predictions broadly followed the 1:1 reference across
treatments, with a modest reduction in scatter for TK under the EC-adjusted model
and substantial overlap between the two models for CK1, CK2, and TC (Figure 3).
Collectively, the blocked validation confirms that cumulative radiation captures the
dominant day-to-day signal, while the additional contribution of EC is scenario
dependent.
Treat
ment
n RMSE
(baselin
e)
MAE
(baseli
ne)
NSE
(baseli
ne)
RMSE
(EC-adjus
ted)
MAE
(EC-adjus
ted)
NSE
(EC-adjusted
)
ΔRMSE
(EC −
baseline)
ΔMAE (EC −
baseline)
ΔNSE
(EC −
baseline
)
CK1 179 1.110 0.905 0.744 1.141 0.926 0.730 0.031 0.021 -0.014
CK2 180 1.020 0.800 0.655 1.032 0.811 0.647 0.012 0.011 -0.008
TK 180 0.829 0.629 0.422 0.815 0.607 0.441 -0.014 -0.022 0.019
TC 182 1.313 0.955 0.473 1.393 0.991 0.407 0.080 0.036 -0.066
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Figure 3. Observed versus predicted daily irrigation volume (I, L d⁻ ¹ per trough) for the blocked
5-fold time-series cross-validation (pooled predictions) under each treatment. Predictions from
both the radiation-only baseline model and the EC-adjusted model are shown. The solid line
indicates the 1:1 reference line.
3.5. Radiation-Driven Irrigation and Drainage Responses
Across treatments, G maintained a highly significant positive association with I
(p < 0.001), and the strength of the relationship varied by scenario. The strongest G–I
correlations occurred under CK1 (r = 0.73) and TC (r = 0.72) (Figure 4), reinforcing
the consistency of radiation-driven water demand under contrasting management
backgrounds.
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Figure 4. Correlation analysis between daily cumulative solar radiation (G, MJ m⁻ ² d⁻¹) and
irrigation/drainage variables (I and D, L d⁻¹ per trough) under different treatments.
Relationships between G and drainage volume were more treatment sensitive.
The G–D correlation was highest under CK2 (r = 0.78), followed by CK1, while
weaker responses were observed under TK and TC (r ≈ 0.51–0.52). This pattern
suggests that drainage reflects not only radiation-driven irrigation input but also
scenario-dependent root uptake and substrate buffering effects.
Correlations between G and solution properties were generally weaker.
Associations with EC irr and pH were mostly small (|r| < 0.35), with significant
negative G–ECirr correlations only under specific treatments. Relationships involving
irrigation and drainage pH were weaker still, indicating that daily variations in EC and
pH were primarily governed by nutrient-solution preparation and substrate buffering
rather than direct radiation forcing.
3.6. Yield Dynamics
Fresh fruit yield per harvest (g plot⁻¹) exhibited pronounced seasonal variation
(Figure 5; S2 Dataset). Low yields in early winter were followed by increasing
harvest amounts toward spring, with peaks in April–May as radiation and temperature
improved.
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Figure 5. Temporal variation in fresh fruit yield (g plot⁻¹ per harvest) under different treatments.
For ‘Jingdan No. 8’ (CK1 and TK), cumulative yield increased steadily
throughout the season. For ‘Jingcai No. 8’ (CK2 and TC), yield remained lower early
in the season and increased later, indicating more stage-dependent responses under the
corresponding water–salt regimes.
Late-season dynamics differed between the two cultivar–scenario groups.
‘Jingcai No. 8’ displayed greater fluctuations: CK2 peaked in mid-April (1.68 kg d⁻¹),
whereas TC peaked in late March (1.43 kg d⁻¹), followed by sharp declines in late
May (CK2: 0.23 kg d⁻¹; TC: 0.12 kg d⁻¹). In contrast, ‘Jingdan No. 8’ maintained
comparatively better late-season performance under both CK1 and TK.
3.7. Fruit Nutritional Quality
Fruit quality was assessed using ascorbic acid/vitamin C (AsA/VC, mg 100 g⁻¹
FW) and sugar–acid ratio (SAR = TSS/TA) under contrasting seasonal radiation
conditions (S3 Dataset), representing low-light winter (January) and high-light spring
(April). Treatment distributions in the AsA/VC–SAR space shifted markedly between
these periods (Figure 6), with lower values in January and higher values in April,
consistent with improved nutritional quality and flavor balance under increased
radiation.
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Figure 6. Fruit quality expressed as ascorbic acid (AsA/VC, mg 100 g⁻ ¹ FW) versus sugar–acid
ratio (SAR = TSS/TA, dimensionless) under different treatments. TSS is in °Brix and titratable
acidity (TA) is in g 100 g⁻¹ FW (citric acid equivalents). The boundary of “high VC + high SAR”
is empirical for reference only.
Within ‘Jingdan No. 8’, CK1 maintained relatively high AsA/VC but lower SAR
than TK. For ‘Jingcai No. 8’, AsA/VC was generally lower across treatments, and TC
remained consistently low. Treatment separation became more apparent under higher
radiation, suggesting that light availability modulated the expression of water–salt
management effects on fruit composition.
3.8. Seasonal Photosynthetic Responses
Seasonal changes in photosynthetic traits are summarized in Figure 7 (S4
Dataset). Net photosynthetic rate (Pn) increased from January to May across
treatments. In January, TK showed the highest mean Pn (13.18 μmol CO₂ m⁻² s ⁻¹),
while TC was lowest (7.99 μmol CO₂ m⁻² s⁻ ¹). In May, mean Pn values converged
across treatments (13.51–14.14 μmol CO₂ m⁻² s⁻¹), with CK1 slightly higher than the
others.
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Figure 7. Temporal variation of photosynthetic parameters (Pn, Gs, Ci, and Tr; units shown on
axes) in tomato under different treatments.
Stomatal conductance (Gs) generally decreased from January to May,
accompanied by a decline in intercellular CO₂ concentration (Ci). Transpiration rate
(Tr) increased seasonally in CK1, CK2, and TC, whereas TK maintained relatively
higher Pn with comparatively lower Tr, indicating scenario-specific decoupling
between carbon assimilation and water loss.
4. Discussion
Limitation
(study design): treatments were cultivar-bound (CK1-TK for 'Jingdan
No. 8'; CK2-TC for 'Jingcai No. 8'), so treatment contrasts are interpreted within each
cultivar-specific case study and we avoid cross-cultivar causal claims.
4.1. Radiation as the Primary Driver of Daily Irrigation Demand
Daily cumulative solar radiation (G) showed consistently strong and highly
significant positive correlations with irrigation volume across all treatments,
confirming radiation-driven transpiration demand as the dominant determinant of
short-term water requirements in soilless greenhouse tomatoes [13,33]. This finding
aligns with previous studies demonstrating that, under protected cultivation, daily
water uptake is more tightly coupled to incident radiation than to air temperature or
humidity alone, particularly when canopy development and root-zone moisture are
maintained within suitable ranges [5,11–14]. Radiation governs stomatal opening, leaf
energy balance, and transpiration flux, thereby directly translating light variability
into irrigation demand at the daily scale [30–32].
Compared with approaches based on stage-based coefficients or
evapotranspiration estimates requiring multiple environmental inputs, the
radiation-driven relationship observed here provides a simplified yet robust basis for
irrigation scheduling [10,15]. Within each cultivar-specific treatment set (CK1–TK
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
for ‘Jingdan No. 8’ and CK2–TC for ‘Jingcai No. 8’), the G–I relationship remained
stable, suggesting that radiation is the primary driver under the management
backgrounds examined. Accordingly, G can be used as the core predictor for daily
irrigation decision-making, while EC and irrigation volume are incorporated as
scenario-specific adjustments rather than as cross-cultivar general rules.
4.2. Scenario-Dependent Effects of Electrical Conductivity on Irrigation Demand
Although radiation dominated daily irrigation variability, the role of irrigation
solution electrical conductivity (EC) exhibited clear scenario dependence. Under
conventional water supply (CK1), EC showed a significant negative association with
irrigation demand, suggesting that higher salinity increased osmotic resistance in the
root zone, thereby suppressing plant water uptake [41]. In contrast, under the
low-water high-EC scenario (TK), EC exerted a positive effect on irrigation demand,
indicating that moderate salinity stress may have enhanced transpiration or induced
compensatory water uptake responses under limited water supply [27–29].
These contrasting responses highlight that EC effects should not be interpreted
independently of irrigation background. Under sufficient water availability, elevated
EC primarily increases osmotic stress and reduces effective water absorption [21,22].
Under restricted irrigation, however, higher EC may alter root hydraulic conductivity,
stomatal regulation, or assimilate partitioning in ways that partially offset water
Limitation
[20, 23]. Similar interaction effects between salinity and irrigation regime
have been reported in greenhouse tomatoes, where moderate salinity improved certain
physiological or quality traits without proportionally reducing water uptake [42].
This scenario dependence also helps interpret why EC improved blocked
cross-validation only under TK. One likely reason is that EC irr was operationally
adjusted in response to radiation (significant negative G–EC irr correlations occurred
only under specific treatments), which can introduce collinearity and temporal
non-stationarity that weakens out-of-sample generalization in CK1/CK2/TC. Under
TK, where irrigation supply is restricted and salinity management is more tightly
coupled to plant water status, EC irr may better reflect osmotic constraints affecting
water uptake and thus provides incremental predictive signal beyond radiation.
Collinearity between G and ECirr was mild in all treatments (pairwise r = -0.26 to
-0.52; VIF = 1.07–1.36), suggesting that the scenario-dependent gain/loss is more
likely driven by how EC was operationally adjusted over time than by severe
multicollinearity.
Consistent with this interpretation, under the high-water moderate-EC scenario
(TC), EC effects on irrigation demand were weak and non-significant, indicating that
abundant water supply can buffer salinity-related impacts on short-term water demand
[8,9]. Together, these results emphasize that EC should be treated as a
scenario-adjustment parameter rather than a universal modifier in radiation-driven
irrigation models.
4.3. Performance and Practical Applicability of Radiation-Driven Empirical Models
The empirical regression models based on daily radiation and EC achieved
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
moderate to high explanatory power (R² = 0.52–0.79) across treatments, with RMSE
values of 0.75–1.13 L d⁻¹ and NSE values closely aligned with R² [16–18,35,36].
These performance metrics indicate that the models effectively captured major
day-to-day fluctuations in irrigation demand, while prediction errors were primarily
associated with occasional extreme deviations rather than systematic bias [13,17].
It is important to emphasize that the proposed models are intended as applied
scheduling tools rather than mechanistic simulations of plant or root-zone processes
[10,34]. By relying on easily measurable inputs and treatment-specific coefficients,
the models balance accuracy with operational feasibility, which is a key consideration
for greenhouse management [11,15]. Similar empirical or semi-empirical approaches
have been widely adopted in protected horticulture to support real-time
decision-making, particularly where full physiological modeling is impractical
[12,16,18].
The blocked cross-validation results further confirm that daily cumulative
radiation is the dominant predictor of day-to-day irrigation demand, as the
radiation-only baseline already provided robust accuracy across treatments. The
additional EC term did not uniformly improve cross-validated errors, but its
contribution was clearly scenario-dependent, consistent with the direction and
significance of EC effects observed in the fitted models. Therefore, EC is better
interpreted as a scenario-adjustment factor for water–salt management rather than a
universal accuracy booster.
4.4. Yield and Quality Responses under Integrated Water–Salt Management
Seasonal yield dynamics reflected the combined influences of radiation
availability, temperature, and water–salt management [2,3,33]. Low yields during
winter coincided with limited radiation, while yield peaks in spring corresponded to
improved light conditions [1,13]. Under the low-water high-EC treatment (TK), yield
accumulation was initially comparable to or slightly higher than the conventional
treatment but declined later in the season, indicating stage-specific sensitivity to
water–salt stress [27,28].
Quality responses showed clearer differentiation among treatments. Higher
ascorbic acid content and sugar–acid ratios under TK, particularly during
high-radiation periods, suggest that moderate water and salinity stress promoted
assimilate concentration and quality enhancement [43]. This trade-off between yield
stability and quality improvement is consistent with previous findings that controlled
stress can enhance fruit quality attributes in greenhouse tomatoes while maintaining
acceptable yield levels [22,23,29,44].
Notably, quality differences among treatments became more pronounced under
higher radiation, indicating that light availability modulates the expression of water–
salt management effects on fruit composition [30–32]. These results underscore the
importance of explicitly integrating radiation conditions when evaluating irrigation
and salinity strategies aimed at balancing yield and quality.
4.5. Variety-Specific Responses, Limitations, and Management Implications
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Varietal responses differed under comparable management frameworks. ‘Jingdan
No. 8’ exhibited greater yield stability and quality potential under both conventional
and low-water high-EC conditions, whereas ‘Jingcai No. 8’ showed weaker
late-season performance under high-water moderate-EC management [2,3,45]. These
differences likely reflect varietal variation in root architecture, salinity tolerance, and
photosynthetic regulation, reinforcing the need for variety-specific optimization rather
than universal irrigation or EC targets [8,9].
Several limitations should be acknowledged. Treatments were variety-bound,
which restricts direct cross-variety comparisons of EC effects. In addition, pH
variation among treatments was limited, preventing robust evaluation of pH–water–
salt interactions [7,39]. Future studies should incorporate factorial designs with
clearer gradients of EC and pH within the same variety, as well as mixed-effects
modeling approaches to better address heterogeneity [16,17].
From a management perspective, the results suggest that radiation-driven
irrigation prediction can serve as the core component of daily scheduling, with EC
used to adjust strategies according to water availability and varietal characteristics
[10,11,15]. Such an approach provides a practical pathway toward integrated water–
salt management in soilless greenhouse tomato production.
5. Conclusions
(1) Daily irrigation demand in soilless greenhouse tomato was primarily driven
by cumulative solar radiation (G), showing a strong positive association with
irrigation amount (I) and drainage (D). A radiation-only linear baseline therefore
provides a parsimonious predictor for day-to-day irrigation forecasting.
(2) In 5-fold blocked time-series cross-validation, the radiation-driven models
achieved RMSE of 0.815–1.393 L d⁻¹ per trough and NSE of 0.407–0.730 across
treatments. Adding irrigation-solution electrical conductivity (EC irr) yielded a small
improvement under the low-water high-EC scenario (TK), but changes were
negligible or negative under CK1, CK2, and the high-water moderate-EC scenario
(TC). Thus, EC should be treated as a scenario-specific adjustment rather than a
universal driver.
(3) Differences in yield, fruit quality, and photosynthetic traits were consistent
with the contrasting water–salt management backgrounds, supporting that irrigation
forecasting should be interpreted together with production targets. However, inference
remains limited to the cultivar–management combinations tested (CK1–TK in
‘Jingdan No. 8’; CK2–TC in ‘Jingcai No. 8’).
(4) For practice, we recommend a two-step scheduling workflow: estimate daily
irrigation from G using the radiation-only baseline, then optionally apply an EC-based
correction where validated (e.g., low-water high-EC conditions), and distribute the
daily total into irrigation events according to greenhouse control settings. This
provides a low-input pathway toward integrated water–salt management for soilless
greenhouse tomato production.
Author Contributions: Conceptualization, Y.Y. and Y.M.; Methodology, L.X., Y.M.
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
and Y.Y.; Formal analysis, L.X. and Q.F.; Investigation, X.G., H.S. and X.L.; Data
curation, L.X. and X.L.; Writing—original draft, L.X.; Writing—review and editing,
Y.Y., Y.M. and Q.F.; Visualization, L.X.; Supervision, Y.Y.; Project administration,
Y.M. and Y.Y.; Funding acquisition, Y.Y.
Funding: This research was supported by the National Key R&D Program of China
(Grant No. 2023YFD2000602); the Major Science and Technology Projects of
Xinjiang Uygur Autonomous Region (Grant Nos. 2022A02005-1, 2022A02005-5);
the Key R&D Program of Xinjiang Uygur Autonomous Region (Grant Nos.
2023B02024-2, 2023B02024-2-1); and the Agricultural Science and Technology
Innovation Steady Support Program of Xinjiang Academy of Agricultural Sciences
(Grant No. xjnkywdzc-2025003-02,xjnkywdzc-2025002-10). The funders had no role
in the study design, data collection and analysis, decision to publish, or preparation of
the manuscript.
Data Availability Statement:All relevant data underlying the findings of this study
are available within the manuscript and its Supporting Information files. Specifically:
S1 Dataset: Daily environmental parameters (radiation, temperature), irrigation
volumes, and drainage records used for modeling.
S2 Dataset: Detailed yield and fruit number data sorted by harvest date.
S3 Dataset: Fruit nutritional quality measurements (Vitamin C, soluble sugar,
titratable acidity).
S4 Dataset: Photosynthetic characteristics (Pn, Gs, Ci, Tr) measured at different
growth stages.
S1 Code: Python script used for analyzing fruit quality and generating Figure 6.
There are no restrictions on data access.
Ethics statement: Not applicable.
Conflicts of Interest: The authors declare no conflict of interest.
References
1. Tang, M.; Lin, R.; Song, J.; Yu, J.; Zhou, Y. Advances in Physiological and Molecular
Mechanisms of Tomato Responses to Light and Temperature Stress. Acta Horticulturae Sinica
2022, 49, 2174–2188.
https://doi.org/10.16420/j.issn.0513-353x.2022-0600.
2. Ma, X.; Chai, X.; Liu, Y.; Li, J. Evaluating the Canopy Light Environment, Photosynthesis, and
Fruit Comprehensive Performance of Greenhouse Tomato under Different Mechanized
Planting Layouts. Horticultural Plant Journal 2025.
3. Zhang, Y.; Li, Y.; Sun, Z.; Li, W.; Liu, X.; Li, T. Estimating the Light Interception and
Photosynthesis of Greenhouse-Cultivated Tomato Crops under Different Canopy
Configurations. Agronomy 2024, 14, 249.
4. Dirlik, I.F.; Kaya, C. Sensor-Guided Smart Irrigation for Tomato Production: Comparing Low
and Optimum Soil Moisture in Greenhouse Environments. Food Energy Security 2025, 14,
e70082.
5. Liu, C.; Gao, H.; Lv, G.; Wu, X.; Zhao, F.; Huo, Z.; Zhang, X. Estimation and Experiment of
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
the Suitable Irrigation Amount of Potted Tomatoes with Coconut Bran Using Pan Evaporation.
Transactions of the Chinese Society of Agricultural Engineering 2022, 38, (issue not
provided).
6. Wang, W.; Zhang, X. Integrated Effects of Irrigation and Fertilization on Yield, Quality and
Water–Fertilizer Use Efficiency of Substrate-Cultured Tomato. Journal of Irrigation and
Drainage 2023, 42, 51–56. https://doi.org/10.13522/j.cnki.ggps.2022218.
7. Song, S.; Lim, R.B.H.; Gaw, L.Y.F.; Richards, D.R.; Tan, H.T.W. Comparison of Vegetable
Production, Resource-Use Efficiency and Environmental Performance of High-Technology
and Conventional Farming Systems for Urban Agriculture. Science of The Total Environment
2022, 807, 150621.
8. Yin, Z.; Cai, J. Combined Effect of Irrigation and Fertilization on Soil Water, Nutrient
Transport, Yield and Water Use Efficiency of Greenhouse Tomato. Journal of Irrigation and
Drainage 2023, 42, 33–44. https://doi.org/10.13522/j.cnki.ggps.2022421.
9. Lu, T.; Wang, H.; Zhang, T.; Shi, C.; Jiang, W. Influence of Electrical Conductivity of Nutrient
Solution in Different Phenological Stages on the Growth and Yield of Cherry Tomato.
Horticulturae 2022, 8, 378.
10. Chen, W.-H.; Yang, F. Semiclosed Greenhouse Climate Control under Uncertainty via
Machine Learning and Data-Driven Robust Model Predictive Control. IEEE Transactions on
Control Systems Technology 2021, 30, 1186–1197.
11. Liu, T.; Han, X.; Hai, Y.; Li, F.; Wang, F.; Xu, W. Construction and Application of a
Weighing Feedback-Based Irrigation System for Facility Tomatoes. Transactions of the
Chinese Society of Agricultural Engineering 2024, 40, 85–96.
12. Xu, D. Effects of Different Aerobic Drip Irrigation Methods on Yield, Quality and
Photosynthesis of Cherry Tomato in Greenhouse. Jiangsu Journal of Agricultural Sciences
2020, 36, 152–157.
13. Shang, C.; Han, Y.; Guo, W. Water Requirement Rule of Tomato in Glasshouse under Rock
Wool Nutrient Solution Cultivation. Journal of Agricultural Science and Technology 2019, 21,
109–117. https://doi.org/10.13304/j.nykjdb.2018.0540.
14. Lozano-Castellanos, L.F.; Lozano-Castellanos, I.C.; Correa-Guimaraes, A. Technologies
Applied to Artificial Lighting in Indoor Agriculture: A Review. Sustainability 2025, 17, 3196.
15. Zhang, Z.; Liu, J.; Kong, T.; Xie, J.; Zhang, M. Design and Experiment of Greenhouse Melon
Irrigation System Based on Environmental VPD Decision. Transactions of the Chinese Society
for Agricultural Machinery 2022, 53, 371–378.
16. Xiang, L.; Li, J. Effects of Irrigation Amount and Frequency on Root Growth, Yield and
Nutrient Absorption of Tomato. Journal of Agricultural Science and Technology of Northwest
A&F University 2024, 52, 80–92.
17. Zhu, C.; Zhang, J.; Xiang, L.; Li, J. Validation of Advanced Decision Irrigation Model of
Greenhouse Tomato Soil Cultivation Based on Environmental Factors. Journal of China
Agricultural University 2023, 28, 60–73.
18. Wang, Z.; Yuan, M.; Yin, W.; Zhang, C.; Zhang, Z.; Hu, X. A Machine Learning-Based
Irrigation Prediction Model for Cherry Tomatoes in Greenhouses. Computers and Electronics
in Agriculture 2025, 237, 110558.
19. Collado, C.; Hernandez, S.; Hernandez, R. Supplemental Greenhouse Lighting Increased the
Water Use Efficiency, Crop Growth, and Cutting Production in Cannabis sativa. Frontiers in
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Plant Science 2024, 15, 1371702.
20. Karaca, C.; Buyuktas, D.; Kurunc, A.; Bastug, R.; Navarro, A. Effects of Salinity Stress on
Drip-Irrigated Tomatoes Grown under Mediterranean-Type Greenhouse Conditions.
Agronomy 2023, 13, 36.
21. Alshami, A.K.; El-Shirbeny, A.; Al-Omran, A.M.; Alghamdi, A.G.; Louki, I.; Alkhasha, A.
Responses of Tomato Crop and Water Productivity to Deficit Irrigation Strategies and Salinity
Stress in Greenhouse. Agronomy 2023, 13, 3016.
22. Pei, S.; Zhang, H.; Zhang, Z.; Zhao, F.; Li, Z. Physiological Response of Potted Tomatoes to
NaCl and Na₂SO₄ Brackish Water Irrigation. Scientia Agricultura Sinica 2024, 57, 570–583.
23. Chen, P.; Dong, X.; Tian, L.; Zhang, X.; Liu, X.; Sun, H. Regulation Effects of Fulvic Acid on
Tomato Yield and Quality under Saline Water Irrigation. Chinese Journal of Eco-Agriculture
2023, 31, 452–462.
https://doi.org/10.12357/cjea.20220178.
24. Linker, R.; Kirzhner, I. Model-Based Simulation–Optimization of Irrigation Scheduling: A
Field Evaluation with Processing Tomatoes. Smart Agricultural Technology 2023, 4, 100234.
25. Liu, H.L., H.; Ning, H.; Zhang, X.; Li, S.; Pang, J.; Wang, G.; Sun, J. Optimizing irrigation
frequency and amount to balance yield, fruit quality and water use efficiency of greenhouse
tomato. Agricultural Water Management, 2019. 226: p. 105787.
26. Incrocci, L.; Malorgio, F.; Pardossi, A. New Trends in the Fertigation Management of
Irrigated Vegetable Crops. Horticulturae 2017, 3, 37.
27. Song, Y.; Zhang, J.; Xing, J.; Wang, L.; Wang, X.; Hu, C.; Li, W.; Tan, Z.; Cheng, Y.
Optimizing Water–Fertilizer Coupling Across Different Growth Stages of Tomato in Yellow
Sand Substrate: Toward Enhanced Yield, Quality and Resource Use Efficiency. Plants 2025,
14, 936.
28. Zhang, P.; Dai, Y. Effects of Salinity Stress at Different Growth Stages on Tomato Growth,
Yield and Water-Use Efficiency. Communications in Soil Science and Plant Analysis 2017,
48, 624–634.
29. Li, J.; Qu, Z.; Wang, S.; He, P.; Zhang, N. Effects of Alternating Irrigation with Fresh and
Saline Water on Soil Salt, Soil Nutrients, and Yield of Tomatoes. Water 2019, 11, 1693.
30. Xiao, L.; Ma, Y.; Wang, X.; Xing, J.; Zheng, J.; Ma, Y. Effects of Supplemental Light Quality
and Substrate Moisture Content on Morphological Regulation and Photosynthetic
Characteristics of Cucumber Seedlings. Journal of Agricultural Science and Technology 2025,
27, 68–77.
https://doi.org/10.13304/j.nykjdb.2023.0733.
31. Marie, T. Growing Tomato in Controlled Environments under Continuous Light Requires
Dynamic LEDs to Entrap the Circadian Rhythm, Adjust Canopy Architecture, and Balance
Photostasis. Ph.D. Thesis, University of Guelph, Guelph, Canada, 2024.
32. He, W.; Li, Z.; Su, W.; Gan, L.; Xu, Z. Effect of Different Light Intensities on the
Photosynthate Distribution in Cherry Tomato Seedlings. The Journal of Horticultural Science
and Biotechnology 2019, 94, 611–619.
33. Hong, M.; Zhang, Z.; Fu, Q.; Liu, Y. Water Requirement of Solar Greenhouse Tomatoes with
Drip Irrigation under Mulch in the Southwest of the Taklimakan Desert. Water 2022, 14, 3050.
34. Sun, L.; Yao, M.; Mao, L.; Zhao, M.; Niu, H.; Xu, Z.; Wang, T.; Wang, J. Simulation of Soil
Water Movement and Root Uptake under Mulched Drip Irrigation of Greenhouse Tomatoes.
Water 2023, 15, 1282.
35. Ge, J.; Gong, X.; Ping, Y.; Luo, J.; Li, Y. Evaluation of Irrigation Modes for Greenhouse Drip
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Irrigation Tomatoes Based on AquaCrop and DSSAT Models. Plants 2023, 12, 3863.
36. Shan, Z.; Zhang, X.; Si, Z.; Yi, R.; Fan, H. Optimizing Irrigation and Nitrogen Application for
Greenhouse Tomato Using the DSSAT–CROPGRO–Tomato Model. Water 2025, 17, 426.
37. Hu, J.; Li, Y.; Hou, J.; Sun, Z.; Wang, H.; He, D. Analysis and Prospect of the Environmental
Control Systems for Greenhouse. Transactions of the Chinese Society of Agricultural
Engineering 2024, 40, 112–128.
38. Liu, X.; Shu, S.; Zhang, A.; Sun, J. Potential Advantages and Theoretical Basis of
Light-Associated Temperature Regulation in Protected Cultivation. Journal of Nanjing
Agricultural University 2023, 46, 823–832.
39. Zhao, W.; Yu, H.; Wang, Z.; Ma, H. Effects and Driving Factors of Water and Fertilizer
Coupling on the Yield and Quality of Tomato Cultivated in Carbon Substrate. Transactions of
the Chinese Society of Agricultural Engineering 2025, 41, 199–207.
40. Savvas, D.; Gruda., N. Application of soilless culture technologies in the modern greenhouse
industry—A review. Eur. J. Hortic. Sci, 2018. 83(5): p. 280-293.
41. Moya, C.; Verdugo, G.; Flores, M.F.; Urrestarazu, M.; Álvaro, J. E.Increased Electrical
Conductivity in Nutrient Solution Management Enhances Dietary and Organoleptic Qualities
in Soilless Culture Tomato. HortScience 2017, 52, 868–872.
42. Bonachela, S.F., M. D.; Cabrera-Corral, F. J.; Granados, M. R. Salt and irrigation management
of soil-grown Mediterranean greenhouse tomato crops drip-irrigated with moderately saline
water. Agricultural Water Management, 2022. 262: p. 107433.
43. Wu, Z.; Qiu, Y.; Hao, X.; Li, S.; Kang, S. Response of Yield and Quality of Greenhouse
Tomatoes to Water and Salt Stresses and Biochar Addition in Northwest China. Agricultural
Water Management 2022, 270, 107736.
44. Roșca, M.M.; Stoleru, V. Tomato Responses to Salinity Stress: From Morphological Traits to
Genetic Changes. Frontiers in Plant Science 2023, 14, 1118383.
45. Shin, J.-Y. Analyses of Canopy Light Interception and Photosynthesis of Greenhouse Tomato
Plants under Diffuse Films Using 3D Plant Model and Ray-Tracing Simulation. Ph.D. Thesis,
Seoul National University, Seoul, Korea, 2020.
.CC-BY 4.0 International licenseperpetuity. It is made available under a
preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
The copyright holder for thisthis version posted January 24, 2026. ; https://doi.org/10.64898/2026.01.23.701235doi: bioRxiv preprint
Text is read by the "Ask this paper" AI Q&A widget below.
Extraction quality varies by source — PMC NXML preserves structure
cleanly, OA-HTML may include some navigation residue, and OA-PDF can
have broken hyphenation. The publisher copy
(via DOI)
is the canonical version.