Physiological and biochemical plasticity as a driver of invasiveness in Syzygium jambos (L.) Alston

Physiological and biochemical plasticity as a driver of invasiveness in Syzygium jambos (L.) 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Alston Cristiano Ferrara Resende, Paulo Henrique Pereira Peixoto This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8196016/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Purpose This study explores the physiological and biochemical plasticity of Syzygium jambos , an invasive tree species native to Tropical Asia, under contrasting light intensities, fertilization regimes, and water availability. The aim is to elucidate the mechanisms underlying its ecological success and to address how these mechanisms may inform strategies for managing invasive plants under changing climate scenarios. Methods A factorial experiment was conducted using seedlings grown in two substrates – control (soil and sand), and organic amendment (aged bovine manure) – and exposed them to three light environments: very low radiation (VLR), low radiation (LR), and high radiation (HR). Experiments were conducted under controlled conditions at the Federal University of Juiz de Fora (Minas Gerais, Brazil). Physiological assessments included gas exchange, chlorophyll a fluorescence, antioxidant enzyme activities, osmotic stress markers, and foliar nutrient profiling. Results Plants under HR exhibited elevated protein content and lipid peroxidation levels, alongside reduced superoxide dismutase (SOD) and polyphenol oxidase (PPO) activities. Organic fertilization enhanced nutrient availability and modulated stress responses. Despite reduced photosynthetic rates under HR, S. jambos maintained photochemical efficiency and water-use balance, with rapid recovery following drought stress. Nutrient analyses revealed significant differences between control and fertilized plants, with fertilized seedlings showing elevated levels of nitrogen (N), phosphorus (P), and potassium (K). Conclusion These findings reveal that S. jambos possesses high ecophysiological plasticity, enabling it to adapt to heterogeneous tropical environments and recover from abiotic stress. Such versatility likely contributes to its invasive potential and ecological dominance. By linking functional traits to invasion success, this study provides a predictive framework for managing exotic species and assessing their impact under climate variability. invasive species photosynthesis chlorophyll a fluorescence water stress light acclimation Figures Figure 1 Figure 2 Figure 3 Introduction Invasive plant species are among the most transformative agents in tropical ecosystems, often reshaping community structure and ecosystem function. Their ecological success is frequently attributed to their ability to tolerate and thrive under a wide range of environmental conditions (Moura et al. 2021 ; Hernández-Fernández et al. 2025 ). Syzygium jambos (L.) Alston (Myrtaceae), commonly known as rose apple, exemplifies this adaptability. Native to Tropical Asia, this fast-growing tree species can exceed 12 meters in height and is widely recognized for its capacity to colonize both disturbed and undisturbed tropical habitats. Its rapid establishment in open areas and mature forests has led to significant alterations in native plant assemblages, particularly through the formation of dense canopies that suppress light-dependent species (Horvitz et al. 1998 ; Aide et al. 2000 ; Lugo 2004 ; Brown et al. 2006 ; Cramer et al. 2008 ; Morales 2020 ). In tropical forests, where light is a limiting and heterogeneously distributed resource, the ability to optimize light use is crucial for plant performance. Light availability drives key physiological processes, including photosynthesis, antioxidant metabolism, and nutrient uptake (Valladares et al. 2000 ; dos Anjos et al. 2015; Liu et al. 2016 ). The interplay between light intensity and soil nutrient availability can significantly shape competitive interactions among species, particularly in environments where resource partitioning is crucial for growth and survival (Ferreira et al. 2009 ). While native species have evolved strategies to cope with such variability, invasive plants often exhibit enhanced phenotypic plasticity, enabling them to outperform native flora under fluctuating conditions (Davidson et al. 2011 ; Chen et al. 2024 ). Invasive plants also tend to exhibit physiological and biochemical traits that enhance their resilience under environmental stress, contributing to their competitive success. Traits such as increased photosynthetic efficiency, flexible stomatal regulation, and robust antioxidant systems enable these species to maintain functional performance across diverse light, nutrient, and water conditions (McAlpine et al. 2008 ; Pintó-Marijuan and Munné-Bosch 2013 ; Sanders et al. 2025 ). Chlorophyll fluorescence is a sensitive tool for assessing photochemical efficiency and has been widely used to assess stress responses in invasive taxa (Zhou et al. 2024 ). Additionally, the activity of antioxidant enzymes – including superoxide dismutase (SOD), catalase (CAT), peroxidase (POD), and polyphenol oxidase (PPO) – plays a central role in mitigating oxidative damage caused by reactive oxygen species (ROS), which tend to accumulate under abiotic stress such as drought, nutrient limitation, and excess light (Wang et al. 2023). These physiological mechanisms, when coupled with phenotypic plasticity, may explain the ability of S. jambos to persist and dominate in diverse tropical environments. Beyond ecological disruption, biological invasions impose substantial economic costs. A global synthesis estimated that the cumulative cost of invasive species reached at least USD 1.28 trillion between 1970 and 2017, with annual costs tripling every decade and peaking at USD 162.7 billion in 2017 alone (Diagne et al. 2021 ). Notably, damage-related expenses – such as loss of ecosystem services and agricultural productivity – far exceed management investments, particularly for plant taxa, which remain underrepresented in economic assessments. In Brazil, the economic burden of biological invasions has recently been quantified for the first time, revealing a minimum cost of USD 105.53 billion over 35 years, with an overwhelming 99% of this amount attributed to damages and losses rather than management efforts (Adelino et al. 2021 ). Given that only 16 invasive species were included in this estimate – out of at least 460 recognized invaders – the actual economic impact is likely much higher. An updated national inventory has since identified 444 invasive non-native species in Brazil, including 188 plants, with trees being the most frequent life form among them (Zenni et al. 2024 ). This reinforces the relevance of investigating arboreal invaders such as S. jambos . These findings underscore the urgency of understanding the physiological mechanisms that underpin invasion success, especially in species like S. jambos , whose ecological dominance is matched by its potential economic impact. This study aimed to investigate the physiological and biochemical traits of S. jambos under varying environmental conditions, in order to identify the features most closely associated with its invasive behavior. By linking ecophysiological performance to invasion potential, we seek to contribute to a more predictive framework for managing exotic species in tropical forest ecosystems. Materials and Methods Plant material and experimental design Fruits of S. jambos were collected from two specimens at the Institute of Biological Sciences (ICB) of the Federal University of Juiz de Fora (UFJF), Minas Gerais, Brazil. Seeds were extracted, placed in Petri dishes with filter paper, and moistened with 20 mL of distilled water to induce imbibition and germination. Germinated seeds were transplanted into 200 mL plastic cups containing Plantmax Hortaliças HT® substrate, kept in shade, and irrigated twice weekly. All seeds developed into seedlings, which were standardized by height (10–12 cm) and leaf count (8–10 leaves) prior to experimental setup. Seedlings were transferred to 25 L pots filled with one of three substrate treatments: 1) control – soil and sand (3:2, v/v); 2) bovine manure – soil, sand, and aged bovine manure (3:2:1, v/v/v); and 3) chemical fertilizer – soil and sand (3:2, v/v) + 50 g of Forth Frutas® (12% N, 5% P, 15% K). The pots were placed in a greenhouse under shade conditions (4% of ambient light, ~ 100 µmol photons m − 2 s − 1 of photosynthetically active radiation – PAR), achieved using Sombrite® screens of different mesh densities. Full sunlight (~ 2,400 µmol photons m − 2 s − 1 of PAR on clear days) was used as the reference for 100% irradiance. Plants remained in this environment for ~ 8 months to allow growth and tissue development. All plants in the chemically fertilized treatment died during this period, leaving only the control group and the bovine manure groups. For the new experimental setup, plants from these two groups were standardized by height (30–50 cm, mean of 40 cm) and the number of leaves (12–18 leaves, mean of 16). They were then assigned to three light treatments: very low radiation (VLR) – 4% of ambient light (~ 100 µmol photons m − 2 s − 1 PAR); low radiation (LR) – 9% of ambient light (~ 220 µmol photons m − 2 s − 1 PAR); and high radiation (HR) – full sunlight (~ 2,400 µmol photons m − 2 s − 1 PAR). The experiments were conducted using a completely randomized factorial design (2 fertilization types × 3 light levels), totaling six treatments. All plants were irrigated at least twice a week and maintained under these conditions for 11 months, after which leaf samples were collected for biochemical analysis. Physicochemical properties and fertility assessment of substrates Prior to the experiment, random samples of the three substrates were collected to assess particle size distribution and nutrient composition. After 11 months of plant growth, three samples from each of the six final treatments were collected and analyzed for their nutritional and chemical attributes. Nutrient extraction followed standard protocols: Mehlich 1 (0.05 M HCl + 0.0125 M H 2 SO 4 ) for K, P, Zn, Fe, Mn, and Cu; 1M KCl for Ca, Mg, and Al; hot water for B; monocalcium phosphate in acetic acid for S; pH in SMP buffer solution for potential acidity (H + Al). Additional parameters included: pH in water (1:2.5), sum of exchangeable bases (SB), cation exchange capacity at pH 7.0 (T), effective cation exchange capacity (t), base saturation index (V), aluminum saturation index (m), organic matter (OM; via oxidation with 4N Na 2 Cr 2 O 7 + 10N H 2 SO 4 ), and remaining phosphorus (P-Rem). Biochemical and enzymatic analysis Crude extracts for total protein and enzymatic activity determinations – superoxide dismutase (SOD; EC 1.15.1.1), catalase (CAT; EC 1.11.1.6), peroxidase (POD; EC 1.11.1.7), and polyphenol oxidase (PPO; EC 1.10.3.1, EC 1.10.3.2, and EC 1.14.18.1) – were obtained by macerating four leaf discs (0.8 cm 2 ; 0.07–0.1 g each) in liquid nitrogen, followed by homogenization in 5 mL of buffer containing 0.1 M potassium phosphate (pH 6.8), 0.1 mM EDTA, and 1 mM PMSF (Peixoto et al. 1999 ). The homogenate was filtered through gauze and centrifuged (10,000 g , 15 min, 4°C). Soluble protein content was determined using the method of Lowry et al. ( 1951 ), with the Folin-Ciocalteu reagent. The protein concentration in each replicate was used to calculate the specific activity of the enzymes. SOD activity was measured according to Del Longo et al. ( 1993 ), with reactions conducted at 25°C under illumination by a 15W fluorescent lamp (Giannopolitis and Ries 1977 ). Photoreduction of nitro blue tetrazolium (NBT) was monitored at 560 nm, and one unit of activity was defined as the amount of enzyme required to inhibit 50% of NBT reduction (Beauchamp and Fridovich 1971 ). CAT activity was measured by monitoring H 2 O 2 decomposition at 240 nm (Havir and McHale 1987 ), using a molar extinction coefficient of 36 M − 1 cm − 1 (Anderson et al. 1995 ). POD and PPO activities were estimated according to Kar and Mishra ( 1976 ), with absorbance measured at 420 nm. A molar extinction coefficient of 2.47 mM − 1 cm − 1 was used for calculations (Chance and Maehley 1954). Proline quantification was performed according to Bates et al. ( 1973 ). Six leaf discs (0.8 cm 2 ; 0.1–0.16 g each) were macerated in liquid nitrogen and homogenized in 10 mL of 3% (w/v) sulfosalicylic acid, then filtered through Whatman No. 2 paper. For each sample, 2 mL of filtrate was combined with 2 mL of acidic ninhydrin and 2 mL of acetic acid, then incubated at 100°C for 1 hour. After cooling in an ice bath, the chromophore was extracted with toluene and separated by decantation. Absorbance was measured at 520 nm, and proline concentration was determined using a standard curve. Hydrogen peroxide (H 2 O 2 ) content was determined using a modified ferrous ammonium sulfate/xylenol orange (FOX) assay, with absorbance measured at 560 nm (Gay and Gebicki 2000 ). Approximately 0.2 g of leaf tissue (7–11 disks, 0.8 cm 2 each) was used, and H 2 O 2 concentration was calculated from a standard curve prepared with authentic H 2 O 2 solutions. Superoxide anion (O 2 − .) content was measured following Mohammadi and Karr ( 2001 ), with modifications. Leaf samples (0.3 g; 40 discs, 0.2 cm² each) were incubated in reaction medium, and superoxide production was quantified based on adrenochrome formation (Misra and Fridovich 1971 ), using a molar extinction coefficient of 4.0 x 10 3 M − 1 cm − 1 (Boveris 1984 ). Lipid peroxidation was assessed by quantifying malondialdehyde (MDA) produced after reaction with thiobarbituric acid (TBA) (Cakmak and Horst 1991 ). Approximately 0.2 g of leaf tissue (25–38 disks, 0.2 cm 2 each) was used per sample. Absorbance was recorded at 532 and 600 nm. MDA-TBA concentration was calculated using a molar extinction coefficient of 155 mM − 1 cm − 1 (Heath and Packer 1968 ). Gas exchange and chlorophyll fluorescence Gas exchange and chlorophyll a fluorescence measurements were conducted after 11 months of plant exposure to different light environments. Analyses were performed between 08:00 and 12:00 h using an infrared gas analyzer (LI-6400XT, Li-Cor, Lincoln, NE, USA) with an LED fluorescence chamber (6400-02B, Li-Cor). Fully expanded, healthy leaves from the fourth or fifth node were selected for use. The leaf chamber was set to a photon flux density of 1,000 µmol photons m − 2 s − 1 and ambient CO 2 concentration (~ 400 µmol mol − 1 ). Leaf temperatures averaged 29 ± 1°C under HR and 34 ± 2°C under VLR and LR. Measured parameters included net photosynthetic rate ( A ), transpiration rate ( E ), stomatal conductance ( g s ), and intercellular CO 2 concentration ( C i ). Derived indices estimated were the ratio of intercellular to external CO 2 concentration ( C i / C a ), water-use efficiency (WUE = A / E ) (Ou et al. 2015 ), intrinsic WUE (WUE int = A / g s ), and carboxylation efficiency ( A / C i ). Fluorescence was assessed on light-adapted leaves exposed to a saturating pulse (6,000 µmol photons m − 2 s − 1 , 0.8 s) to determine steady-state fluorescence (F s ) and maximum fluorescence under light (F m ’). Far-red light was then applied to measure the minimum fluorescence of light-adapted leaves (F o ’). Based on Genty et al. ( 1989 ), the following parameters were calculated: effective quantum yield of PSII electron transport (Φ PSII = (F m ’ – F s )/F m ’), efficiency of excitation energy capture by open PSII reaction centers (F v ’/F m ’ = (F m ’ – F o ’)/F m ’), the photochemical quenching (q P = (F m ’ – F s )/(F m ’ – F o ’)), and the apparent electron transport rate (ETR = Φ PSII × PAR × 0.5 × 0.84, where PAR is the photosynthetically active radiation incident on the leaf, 0.5 represents the fraction of photons allocated to photosystem I and photosystem II, and 0.84 is the average fraction of incident light absorbed by the leaf). Leaf nutrient contents Macro- and micronutrient contents in leaves were determined at the end of the experiments. Samples were dried at 65°C to constant weight, ground, and analyzed using standardizing methodologies. The nutrients evaluated included N, P, K, Ca, Mg, S, B, Cu, Mn, Zn, and Fe. Nitrogen (N) content was determined using the sulfuric digestion method followed by quantification via the Kjeldahl procedure. For K, S, Fe, Ca, Zn, Mg, Cu, and Mn, tissue samples were digested in a nitric-perchloric solution (3:1; v:v). Boron (B) was extracted using a hydrochloric acid (HCl) solution. Nutrient concentrations were determined by atomic absorption spectrophotometry (for K, Ca, Mg, Fe, Cu, Mn, and Zn) and colorimetric spectrophotometry (for P, S, and B). Drought stress experiment After 11 months of growth, plants from the control group under the two most contrasting light conditions – VLR and HR – were selected for a drought stress experiment. These plants were subjected to a progressive dehydration protocol, in which irrigation was withheld for 10 or 20 days, while a well-watered control group continued to receive daily irrigation. This design resulted in six distinct treatments. Following the drought phase, all plants were rehydrated and maintained under optimal watering conditions. Leaf samples were collected at the end of each drought period (10 or 20 days without irrigation) and one week after rehydration for physiological assessments. These included measurements of gas exchange parameters and antioxidant enzyme activities, as previously described. Soil moisture and temperature were continuously monitored throughout the experiment. Predawn leaf water potential was determined during both dehydration and rehydration phases using a Scholander-type pressure chamber, as described by Turner ( 1988 ). Statistical analysis Data were first tested for homogeneity of variances (Cochran-Bartlett test) and normality (Lilliefors). Once these assumptions were met, data were subjected to analysis of variance (ANOVA), and means were grouped using the Scott-Knott test at a 5% probability level, performed with SAEG software (version 9.1). Results Chemical analyses of substrates Before the cultivation, the substrate pH was near neutral in the control group (6.9) and slightly acidic in the bovine manure-fertilized group (6.2). In contrast, the chemically fertilized substrate was much more acidic (pH 4.5) (Table 1 ). Potential acidity (H + Al) was also markedly higher in the chemically fertilized material – 4.5 times greater than the control and 3.6 times greater than the manure-fertilized group. Macronutrient (K, P, Ca, Mg, S) and micronutrient (Mn, B) levels were highest in the chemically fertilized substrate, followed by the manure-fertilized and control groups. Fe and Cu contents decreased in the order: chemically fertilized, control, and manure fertilized. Zn and organic matter (OM) were most abundant in the manure-fertilized substrate. Despite differences in P levels, P-Rem and base saturation (V) were similar across treatments. Confirmed by the aluminum saturation index (m), Al was detected only in the chemically fertilized substrate. Although classified as different soil types – medium texture (control and chemical) and clayey texture (manure) – the proportions of clay, silt, and sand were similar among them, with sand predominating. Table 1 Chemical characterization and granulometry of substrates used in the experiment. Parameters: pH of the substrates (pH); contents of potassium (K), phosphorus (P), calcium (Ca), magnesium (Mg), aluminum (Al), zinc (Zn), iron (Fe), manganese (Mn), copper (Cu), boron (B), sulfur (S), remaining phosphorus (P-Rem), organic matter (OM); potential acidity (H + Al); sum of exchangeable bases (SB), effective cation exchange capacity (CEC) (t), and CEC at pH 7.0 (T); base saturation index (V) and aluminum saturation index (m); clay, silt, and sand. Soil type 2 – medium texture; soil type 3 – clayey texture Parameter Substrate Control Chemical fertilization Bovine manure pH 6.9 4.5 6.2 K ( mg dm − 3 ) 30.00 720.00 396.00 P ( mg dm − 3 ) 10.24 121.49 18.58 Ca (cmol dm − 3 ) 4.10 12.71 5.30 Mg (cmol dm − 3 ) 1.00 4.58 2.20 Al (cmol dm − 3 ) 0.00 0.20 0.00 Zn (mg dm − 3 ) 26.41 58.29 102.24 Fe (mg dm − 3 ) 82.67 96.36 66.68 Mn (mg dm − 3 ) 33.22 67.55 59.86 Cu (mg dm − 3 ) 0.77 5.96 0.68 B (mg dm − 3 ) 0.17 0.84 0.50 S (mg dm − 3 ) 37.91 135.64 91.28 P-Rem (mg L − 1 ) 15.94 12.93 14.17 O.M (dag kg − 1 ) 1.87 2.61 3.84 H + Al (cmolc dm − 3 ) 1.55 7.04 1.94 SB (cmolc dm − 3 ) 5.18 19.14 8.52 t (cmolc dm − 3 ) 5.18 19.34 8.52 T (cmolc dm − 3 ) 6.73 26.18 10.46 V (%) 76.92 73.09 81.41 m (%) 0.00 1.03 0.00 Clay (dag kg − 1 ) 28 34 37 Silt (dag kg − 1 ) 3 5 4 Sand (dag kg − 1 ) 69 61 59 Classification Type 2 Type 2 Type 3 Since plants treated with chemical fertilizer died before the start of light intensity treatments, only the control and bovine manure-fertilized substrates were evaluated. Performed 11 months after plant development, nutritional analyses of the substrates showed that in the control group, radiation intensity had no significant effect on most soil parameters, except for S levels, which decreased with increasing light intensity (Table 2 ). In contrast, bovine manure-fertilized substrates showed marked differences in the concentration of essential elements across light environments. After nearly one year of cultivation, substrate pH under HR was significantly higher (mean 7.03) than in LR and VLR. Under VLR, levels of K, P, Mg, B, sum of exchangeable bases (SB), and cation exchange capacity (CEC) at pH 7.0 (T) were significantly more elevated, with no differences between LR and HR. Ca, Zn, Fe, Mn, remaining phosphorus (P-Rem), organic matter (OM), and base saturation index (V) showed no significant variation across light treatments. Cu was lower under HR, and potential acidity (H + Al) was reduced under VLR. S levels mirrored those in the control, decreasing with higher radiation. Al was not detected in any of the samples. No significant differences in pH or in the Zn, Fe, Cu, and S concentrations were found between control and bovine manure-fertilized substrates across light environments. After 11 months, substrates prepared with manure generally showed higher levels of K, P, Ca, Mg, P-Rem, OM, SB, T, and V. Compared to the control, higher concentrations of Mn and B were detected only in the fertilized material under VLR. In VLR and LR conditions, H + Al levels were higher in control than in those with organic amendment. Table 2 Chemical attributes of substrates of Syzygium jambos plants after 11 months of development in three light environments (very low radiation – VLR, low radiation – LR, and high radiation – HR), subjected to two fertilization systems (control group and bovine manure-fertilized). Parameters: pH of the substrates (pH); contents of potassium (K), phosphorus (P), calcium (Ca), magnesium (Mg), zinc (Zn), iron (Fe), manganese (Mn), copper (Cu), boron (B), sulfur (S), remaining phosphorus (P-Rem), organic matter (OM); potential acidity (H + Al); sum of exchangeable bases (SB), cation exchange capacity (CEC) at pH 7.0 (T); and base saturation index (V). Uppercase letters compare the effect of radiation within each fertilization system, while lowercase letters compare the effect of fertilization within each light intensity. Data are presented as mean ± standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability Parameter Control Bovine manure VLR LR HR VLR LR HR pH 6.60 ± 0.06 Aa 6.87 ± 0.33 Aa 6.80 ± 0.06 Aa 6.37 ± 0.30 Ba 6.70 ± 0.10 Ba 7.03 ± 0.03 Aa K ( mg dm − 3 ) 26.67 ± 1.76 Ab 26.00 ± 0.00 Ab 25.33 ± 2.40 Ab 199.33 ± 37.81 Aa 110.00 ± 12.49 Ba 64.67 ± 6.77 Ba P ( mg dm − 3 ) 10.25 ± 0.46 Ab 9.08 ± 0.66 Ab 8.44 ± 0.50 Ab 21.37 ± 2.01 Aa 16.83 ± 0.17 Ba 16.02 ± 0.76 Ba Ca (cmolc dm − 3 ) 4.13 ± 0.15 Ab 3.93 ± 0.13 Ab 3.97 ± 0.09 Aa 4.87 ± 0.37 Aa 4.53 ± 0.13 Aa 4.23 ± 0.13 Aa Mg (cmolc dm − 3 ) 1.10 ± 0.06 Ab 1.00 ± 0.06 Ab 0.97 ± 0.03 Ab 2. 07 ± 0.18 Aa 1.77 ± 0.07 Ba 1.57 ± 0.07 Ba Zn (mg dm − 3 ) 24.73 ± 1.09 Aa 22.08 ± 2.82 Aa 26.51 ± 1.32 Aa 24.64 ± 1.98 Aa 26.35 ± 0.88 Aa 22.38 ± 2.88 Aa Fe (mg dm − 3 ) 54.52 ± 0.90 Aa 60.00 ± 5.25 Aa 64.85 ± 12.98 Aa 69.93 ± 2.64 Aa 69.93 ± 5.88 Aa 65.70 ± 5.54 Aa Mn (mg dm − 3 ) 28.20 ± 2.75 Ab 32.49 ± 2.93 Aa 32.74 ± 1.88 Aa 43.16 ± 5.03 Aa 43.45 ± 2.24 Aa 42.02 ± 6.51 Aa Cu (mg dm − 3 ) 0.64 ± 0.06 Aa 0.57 ± 0.07 Aa 0.56 ± 0.03 Aa 0.68 ± 0.03 Aa 0.68 ± 0.02 Aa 0.52 ± 0.07 Ba B (mg dm − 3 ) 0.14 ± 0.01 Ab 0.19 ± 0.01 Aa 0.15 ± 0.02 Aa 0.23 ± 0.04 Aa 0.16 ± 0.01 Ba 0.14 ± 0.01 Ba S (mg dm − 3 ) 57.47 ± 5.76 Aa 43.18 ± 6.99 Ba 19.24 ± 0.98 Ca 68.05 ± 5.99 Aa 31.66 ± 1.50 Ba 16.90 ± 1.85 Ca P-Rem (mg L − 1 ) 14.65 ± 1.00 Ab 14.75 ± 0.29 Ab 13.34 ± 0.41 Ab 18.04 ± 0.60 Aa 17.05 ± 0.43 Aa 17.37 ± 0.28 Aa OM (dag kg − 1 ) 2.11 ± 0.00 Ab 2.19 ± 0.08 Ab 2.07 ± 0.04 Ab 2.53 ± 0.08 Aa 2.61 ± 0.15 Aa 2.79 ± 0.21 Aa H + Al (cmolc dm − 3 ) 1.82 ± 0.08 Aa 1.89 ± 0.08 Aa 1.65 ± 0.09 Aa 1.36 ± 0.06 Bb 1.60 ± 0.04 Ab 1.68 ± 0.13 Aa SB (cmolc dm − 3 ) 5.30 ± 0.21 Ab 5.00 ± 0.19 Ab 5.00 ± 0.12 Aa 7.44 ± 0.63 Aa 6.58 ± 0.22 Ba 5.96 ± 0.22 Ba T (cmolc dm − 3 ) 7.12 ± 0.20 Ab 6.89 ± 0.18 Ab 6.65 ± 0.17 Ab 9.14 ± 0.59 Aa 8.18 ± 0.26 Ba 7.65 ± 0.16 Ba V (%) 74.39 ± 1.24 Ab 72.51 ± 1.24 Ab 75.14 ± 1.05 Aa 81.40 ± 3.29 Aa 80.42 ± 0.17 Aa 77.99 ± 1.80 Aa Comparisons between initial and post-cultivation substrate fertility revealed notable shifts (Tables 1 and 2 ). pH decreased in unfertilized substrates but increased in those with bovine manure, mirroring trends in potential acidity (H + Al). Macronutrients (K, P, Ca, Mg, S) and parameters SB, t, and T declined more sharply in manure-amended substrates, proportionally to light intensity. Micronutrient levels (Zn, Fe, Mn, Cu, B) fluctuated without consistent patterns. After 11 months, OM increased in control but declined in substrates with manure addition, while P-Rem showed the opposite trend. Foliar nutrient concentrations The results for foliar nutrient concentrations are summarized in Table 3 . In the control group, the concentrations of P, K, Ca, Mg, B, Mn, and Fe did not vary significantly with light intensity. However, foliar N decreased as light radiation increased, with significant differences observed in both control and manure-fertilized plants. In control plants, S was highest under VLR, while Cu and Zn peaked under HR. In manure-fertilized plants, most nutrients varied significantly across light treatments, except for Fe, which remained stable. HR conditions favored the accumulation of P, Mg, B, Cu, and Zn, while the opposite was observed for K, Ca, and S (higher concentrations in VLR). Across all light environments, S, Zn, and Fe showed no significant differences between control and fertilized groups. In general, fertilized plants showed higher levels of N, P, and K, although these differences were not always statistically significant. Notably, B and Cu were significantly higher in fertilized plants under HR. Control plants had higher Ca and Mg overall, except for Mg under HR. Table 3 Foliar nutrient concentrations in Syzygium jambos plants after 11 months of growth under three light environments (very low radiation – VLR; low radiation – LR; and high radiation – HR) and two fertilization systems (control group and bovine manure-fertilized). Uppercase letters compare the effect of radiation within each fertilization system, while lowercase letters compare the effect of fertilization within each light intensity. Data are presented as mean ± standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability Parameter Control Bovine manure VLR LR HR VLR LR HR N (%) 1.39 ± 0.04 Ab 0.99 ± 0.07 Bb 0.78 ± 0.04 Cb 1.67 ± 0.03 Aa 1.23 ± 0.06 Ba 0.97 ± 0.06 Ca P (%) 0.08 ± 0.01 Aa 0.06 ± 0.00 Ab 0.06 ± 0.01 Ab 0.09 ± 0.01 Ba 0.10 ± 0.01 Ba 0.13 ± 0.02 Aa K (%) 0.63 ± 0.04 Ab 0.52 ± 0.05 Ab 0.37 ± 0.02 Aa 1.33 ± 0.12 Aa 0.76 ± 0.04 Ba 0.54 ± 0.09 Ca Ca (%) 0.88 ± 0.02 Aa 0.84 ± 0.03 Aa 0.77 ± 0.03 Aa 0.70 ± 0.03 Ab 0.56 ± 0.05 Bb 0.57 ± 0.02 Bb Mg (%) 0.36 ± 0.02 Aa 0.35 ± 0.00 Aa 0.36 ± 0.01 Aa 0.27 ± 0.01 Bb 0.29 ± 0.01 Bb 0.33 ± 0.02 Aa S (%) 0.17 ± 0.01 Aa 0.13 ± 0.00 Ba 0.11 ± 0.01 Ba 0.17 ± 0.01 Aa 0.14 ± 0.00 Ba 0.13 ± 0.00 Ba B (ppm) 15.3 ± 2.5 Aa 11.7 ± 1.1 Aa 24.3 ± 6.8 Ab 19.4 ± 0.7 Ba 22.3 ± 4.5 Ba 35.8 ± 3.0 Aa Cu (ppm) 0.53 ± 0.24 Ba 0.37 ± 0.12 Ba 1.33 ± 0.38 Ab 0.87 ± 0.20 Ba 1.03 ± 0.23 Ba 2.33 ± 0.28 Aa Mn (ppm) 48.2 ± 35.2 Aa 14.8 ± 3.1 Ab 6.1 ± 2.8 Aa 10.7 ± 3.5 Ba 167.9 ± 76.6 Aa 21.5 ± 9.5 Ba Zn (ppm) 13.1 ± 0.9 Ba 13.0 ± 0.3 Ba 20.4 ± 1.5 Aa 12.4 ± 0.5 Ba 11.3 ± 0.3 Ba 21.3 ± 1.2 Aa Fe (ppm) 131.3 ± 19.3 Aa 131.3 ± 38.5 Aa 88.4 ± 4.0 Aa 129.7 ± 5.5 Aa 110.2 ± 6.2 Aa 93.6 ± 5.2 Aa Biochemical analyses Regardless of the fertilization method or measurement unit, plants under HR had significantly higher total protein content (Table 4 ). In the control group, proline accumulated more under HR, while in fertilized plants, it was higher under VLR. SOD and PPO activities were lowest under HR, especially in fertilized plants, which showed significant variation across light treatments. CAT activity remained unchanged in fertilized plants but decreased under LR in the control group, whereas POD activity peaked under LR in both groups. Overall, hydrogen peroxide (H 2 O 2 ) and superoxide (O 2 . − ) levels were lowest under HR, while MDA content was highest under HR, regardless of fertilization. Comparing control and fertilized plants, protein content differed significantly only under HR, with higher values in the control group. Proline was higher in fertilized plants under VLR and in control plants under HR. SOD activity differed only under LR, being lower in fertilized plants. CAT activity varied under VLR and HR, with fertilized plants showing the lowest values. POD and PPO activities differed only under VLR, with higher POD in control and higher PPO in fertilized plants. Hydrogen peroxide, superoxide, and MDA levels were generally similar between groups, except for MDA under HR, which was higher in fertilized plants. Table 4 Biochemical attributes of Syzygium jambos plants after 11 months of growth under three light environments (very low radiation – VLR, low radiation – LR, and high radiation – HR) and two fertilization systems (control group and bovine manure-fertilized). Total protein content is presented on a fresh mass basis (mg g − 1 FM) and area basis (mg cm − 2 ); proline content (µmol g − 1 FM); enzymatic activities of SOD (units mg − 1 protein), CAT (µmol of H 2 O 2 min − 1 mg − 1 protein), POD (µmol mg − 1 protein), and PPO (µmol mg − 1 protein); and contents of peroxide (nmol g − 1 FM), superoxide (nmol g − 1 FM), and malondialdehyde (MDA; µmol g − 1 FM). Uppercase letters compare the effect of radiation within each fertilization system, while lowercase letters compare the effect of fertilization within each light intensity. Data are presented as mean ± standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability. Parameter Control Bovine manure VLR LR HR VLR LR HR Proteins (/mass) 26.0 ± 1.8 Ba 23.9 ± 1.4 Ba 93.6 ± 2.5 Aa 20.3 ± 0.9 Ba 20.8 ± 0.3 Ba 81.8 ± 3.5 Ab Proteins (/area) 0.60 ± 0.03 Ba 0.60 ± 0.04 Ba 3.23 ± 0.11 Aa 0.49 ± 0.02 Ba 0.53 ± 0.01 Ba 2.72 ± 0.16 Ab Proline 0.54 ± 0.04 Bb 0.34 ± 0.05 Ba 0.82 ± 0.07 Aa 0.75 ± 0.12 Aa 0.43 ± 0.04 Ba 0.54 ± 0.07 Bb SOD 24.47 ± 0.37 Aa 22.40 ± 0.95 Aa 1.17 ± 0.21 Ba 24.82 ± 2.35 Aa 18.05 ± 1.05 Bb 0.02 ± 0.01 Ca CAT 2.81 ± 0.41 Aa 0.49 ± 0.31 Ba 2.91 ± 0.13 Aa 1.20 ± 0.38 Ab 0.49 ± 0.31 Aa 0.95 ± 0.13 Ab POD 0.39 ± 0.04 Ba 0.60 ± 0.07 Aa 0.11 ± 0.01 Ca 0.08 ± 0.04 Bb 0.62 ± 0.10 Aa 0.16 ± 0.01 Ba PPO 1.46 ± 0.22 Ab 1.75 ± 0.09 Aa 0.29 ± 0.01 Ba 2.27 ± 0.13 Aa 1.54 ± 0.09 Ba 0.28 ± 0.04 Ca Peroxide 70.3 ± 17.1 Aa 65.5 ± 6.0 Aa 29.7 ± 3.2 Ba 53.7 ± 12.7 Aa 43.0 ± 1.5 Aa 27.4 ± 2.8 Aa Superoxide 22.50 ± 6.30 Aa 21.41 ± 3.47 Aa 3.73 ± 1.47 Ba 32.20 ± 2.92 Aa 15.52 ± 4.26 Ba 6.67 ± 1.38 Ba MDA 1.25 ± 0.05 Ba 1.31 ± 0.02 Ba 1.54 ± 0.07 Ab 1.34 ± 0.06 Ba 1.39 ± 0.01 Ba 1.89 ± 0.07 Aa Gas exchange and chlorophyll a responses Higher light intensities significantly reduced net CO 2 assimilation ( A ), transpiration ( E ), stomatal conductance ( g s ), and carboxylation efficiency ( A / C i ) (Fig. 1 ). WUE increased with light intensity across all treatments, regardless of fertilization. In the control group, WUE int followed the same pattern, while fertilized plants showed the opposite trend, with a peak under VLR. Fertilized plants showed reduced A , E , g s , and C i compared to controls, except for A under VLR and for E and C i under HR, where differences were not significant. WUE and WUE int were higher in fertilized plants under VLR but declined under HR, reversing the initial advantage. A / C i remained higher in fertilized plants under VLR, while under HR, control plants surpassed them. Figure 2 presents the fluorescence parameters across treatments. Overall, F v ’/F m ’ was lower in plants under HR compared to other light conditions. Φ PSII and ETR followed a similar pattern, with the highest values under VLR, regardless of fertilization. Comparing control and manure-fertilized plants, F v ’/F m ’ was significantly higher in the control group, while Φ PSII , q p , and ETR were consistently higher in fertilized plants across light intensities. The exception was under HR, where Φ PSII and ETR showed no significant differences between groups. Gas exchanges under dehydration and after rehydration The most pronounced changes in gas exchange occurred across dehydration durations within each light environment (Table 5 ). Under VLR, A remained stable after 10 days but declined significantly after 20 days. Under HR, A dropped more sharply, especially after 20 days. E followed a similar trend, with greater reductions under HR. g s decreased significantly in both light conditions after 20 days, with a more pronounced effect under HR. WUE and WUE int means varied with dehydration time but showed no significant differences between treatments. A / C i remained unchanged, except after 20 days in plants under HR, when values were lower. Following rehydration, differences among dehydration treatments diminished, and values of A , E , g s , and A / C i increased, particularly under HR, where the recovery was statistically significant. During the experiment, a significant decline in soil moisture was observed after 10 and 20 days of dehydration under both VLR and HR conditions (Table 6 ). Plant water potential also decreased with dehydration, particularly under HR, dropping from − 0.20 MPa in the control to -4.63 MPa after 20 days (Fig. 3 ). Following rehydration, plant water potential showed marked recovery, with no significant differences among treatments. Comparing plants subjected to 20 days of dehydration before and after rehydration, the increase in water potential was significant under HR. Table 5 Gas exchange parameters in Syzygium jambos plants subjected to different dehydration durations (10 and 20 days) under very low radiation (VLR) and high radiation (HR), before and after rehydration. Parameters include CO 2 assimilation rate ( A , µmol CO 2 m − 2 s − 1 ), transpiration rate ( E , mmol H 2 O m − 2 s − 1 ), stomatal conductance ( g s , mol H 2 O m − 2 s − 1 ), intracellular-to-ambient CO 2 ratio ( C i / C a ), water-use efficiency (WUE, µmol CO 2 mmol − 1 H 2 O), intrinsic water use efficiency (WUE int , µmol CO 2 mol − 1 H 2 O), and carboxylation efficiency ( A / C i , µmol m − 2 s − 1 Pa − 1 ). Uppercase letters indicate comparisons of dehydration duration within each light condition, while lowercase letters compare the effects of light conditions within each dehydration treatment. Data are presented as mean ± standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability. Asterisks denote statistically significant differences between dehydration and post-rehydration values (p < 0.05) Under dehydration VLR HR Parameter control 10 days 20 days control 10 days 20 days A 5.04 ± 0.10 Aa 4.45 ± 0.45 Aa 0.60 ± 0.18 Ba 5.09 ± 0.30 Aa 2.35 ± 0.17 Bb 0.12 ± 0.01 Ca E 1.19 ± 0.11 Aa 0.98 ± 0.10 Aa 0.07 ± 0.00 Ba 0.98 ± 0.03 Aa 0.38 ± 0.07 Bb 0.02 ± 0.00 Ca g s 0.097 ± 0.013 Aa 0.072 ± 0.010 Aa 0.004 ± 0.000 Ba 0.050 ± 0.002 Ab 0.018 ± 0.004 ABb 0.001 ± 0.000 Ba C i / C a 0.76 ± 0.03 Aa 0.72 ± 0.03 ABa 0.33 ± 0.16 Ba 0.56 ± 0.01 Aa 0.42 ± 0.07 Aa 0.36 ± 0.12 Aa WUE 4.33 ± 0.45 Aa 4.59 ± 0.52 Aa 8.59 ± 2.19 Aa 5.21 ± 0.13 Aa 6.49 ± 0.76 Aa 7.18 ± 1.55 Aa WUE int 54.0 ± 7.3 Aa 63.6 ± 7.5 Aa 162.1 ± 41.0 Aa 101.8 ± 1.8 Aa 137.6 ± 17.0 Aa 153.9 ± 31.4 Aa A / C i 0.017 ± 0.001 Aa 0.016 ± 0.002 Aa 0.008 ± 0.005 Aa 0.023 ± 0.002 Aa 0.015 ± 0.002 Aa 0.001 ± 0.000 Ba After rehydration A 4.90 ± 0.23 Aa 4.86 ± 0.33 Aa 3.68 ± 0.62 Aa* 4.81 ± 0.40 Aa 3.18 ± 0.28 Aa 3.66 ± 0.44 Aa* E 1.18 ± 0.11 Aa 1.08 ± 0.04 Aa 0.73 ± 0.16 Aa 1.07 ± 0.20 Aa 0.61 ± 0.11 Aa 0.76 ± 0.06 Aa* g s 0.097 ± 0.013 Aa 0.063 ± 0.004 ABa 0.042 ± 0.010 Ba 0.075 ± 0.018 Aa 0.032 ± 0.006 Aa 0.040 ± 0.003 Aa* C i / C a 0.77 ± 0.03 Aa 0.66 ± 0.01 Aa 0.59 ± 0.06 Aa 0.70 ± 0.04 Aa 0.55 ± 0.06 Aa 0.61 ± 0.02 Aa WUE 4.24 ± 0.52 Aa 4.50 ± 0.16 Aa 5.27 ± 0.53 Aa 4.71 ± 0.53 Aa 5.48 ± 0.73 Aa 4.81 ± 0.21 Aa WUE int 52.9 ± 8.2 Aa 77.3 ± 1.6 Aa 94.9 ± 14.1 Aa 69.5 ± 10.9 Aa 105.1 ± 13.9 Aa 91.5 ± 5.4 Aa A / C i 0.016 ± 0.001 Aa 0.018 ± 0.001 Aa 0.016 ± 0.002 Aa 0.017 ± 0.001 Aa 0.015 ± 0.001 Aa 0.015 ± 0.002 Aa* Table 6 Soil temperature, soil moisture, and leaf water potential (Ψw) of Syzygium jambos plants subjected to dehydration for 10 and 20 days under very low radiation (VLR) and high radiation (HR), followed by rehydration. Uppercase letters indicate comparisons of dehydration duration within each light condition, while lowercase letters compare the effects of light conditions within each dehydration treatment. Data are presented as mean ± standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability. Asterisks denote statistically significant differences between dehydration and post-rehydration values (p < 0.05) Under dehydration VLR HR Parameter control 10 days 20 days control 10 days 20 days Soil temperature (°C) 21.8 ± 0.0 Ab 25.5 ± 0.1 Ab 25.8 ± 0.6 Aa 27.4 ± 1.2 Ba 33.5 ± 1.2 Aa 29.8 ± 1.5 ABa Soil moisture (%) 8.1 ± 0.4 Aa 0.7 ± 0.1 Ba 0.7 ± 0.1 Ba 3.5 ± 0.5 Ab 0.6 ± 0.0 Ba 0.6 ± 0.1 Ba Ψw (MPa) -0.36 ± 0.03 Aa -0.51 ± 0.05 Aa -1.99 ± 0.54 Ba -0.20 ± 0.01 Aa -1.80 ± 0.25 Bb -4.63 ± 0.09 Cb After rehydration Ψw (MPa) -0.36 ± 0.03 Aa -0.54 ± 0.10 Aa -0.32 ± 0.06 Aa -0.21 ± 0.04 Aa -0.38 ± 0.11 Aa -0.20 ± 0.01 Aa* Discussion Soil pH, or active acidity, reflects H + concentration in solutions. Based on classification of Soil Survey Staff ( 2024 ), the control treatment had pH within the “neutral” range (6.6–7.3), the bovine manure treatment had pH values within the “slightly acid” range (6.1–6.5), with the manure-amended substrate within the ideal range for plant growth (5.5–6.5), at pH 6.2 (Islam et al. 1980 ). The chemically fertilized substrate falls within the “very strong acid” range (4.5-5.0), nearing the “extremely high” threshold (3.5–4.4). Excess acidity can increase the solubility of toxic elements, such as manganese and aluminum, which impair root function and cell development (Kochian 1995 ; Horst et al. 2010 ). This pH condition can potentially reduce yield, which may have been the primary factor in the death of plants observed in treatments supplemented with chemical fertilizer. Organic amendment (with bovine manure) enriched the substrates with nutrients and improved chemical attributes by increasing pH and base saturation while reducing acidity. The absence of Al indicates that the amendment effectively buffered soil acidity, preventing Al mobilization. Over the course of the experiment, pH values became more basic in fertilized substrates. This shift is likely linked to the decomposition of manure, which releases carbonates, bicarbonates, and organic acids – compounds that buffer acidity and raise substrate pH (Whalen et al. 2000 ; Holatko et al. 2022 ). Before planting, manure-amended substrates contained higher levels of most macro- and micronutrients than the control (except Fe and Cu). This pattern largely persisted after cultivation for K, P, Ca, Mg, OM, SB, t, T, and V across all light environments. Thus, while manure improved substrate fertility and buffering capacity, the intensity of radiation determined the rate and direction of nutrient transformations in the soil. Over the 11-month experimental period, nutrient depletion was more pronounced in manure-amended substrates under HR, especially for K, P, Mg, Cu, B and S, as well as for the parameters SB and T, reflecting greater nutrient demand for plant growth. Foliar N content decreased with increasing light intensity in both control and fertilized plants, consistent with findings in Sinarundinaria nitida , where shaded plants maintained higher N and chlorophyll levels (Yang et al. 2014b ). In the control group, P, K, Ca, Mg, B, and Mn remained relatively stable across light environments, while in the manure-treated group, nutrient levels fluctuated without a clear pattern. Fertilized plants accumulated, under high radiation, more P, Mg, Cu, B, and Zn, whereas K, Ca, and S decreased. These variable responses align with reports of light-dependent nutrient allocation in other tropical tree species, such as Swietenia macrophylla and Dipteryx odorata (Gonçalves et al. 2005 ), and with studies noting inconsistent effects of light intensity on macronutrient composition (Ronquim et al. 2009 ; Dalmolin et al. 2012 ). These patterns in nutrient dynamics also help explain the observed physiological responses. Plants under HR exhibited higher total protein content than those under VLR and LR, regardless of fertilization. Interestingly, despite lower foliar N levels under HR, protein content was higher. This apparent discrepancy reflects the complex distribution of nitrogen within leaf tissues, which can shift between photosynthetic and structural pools depending on environmental conditions (Wang et al. 2021 ). According to these authors, foliar nitrogen is allocated among several functional compartments, including photosynthetic proteins such as RuBisCo and components of the light-harvesting complexes, as well as non-photosynthetic proteins in cell walls, mitochondria, and cytosol. S. jambos also exhibited higher chlorophyll concentrations under VLR, as confirmed by spectrophotometric analysis and by non-destructive analyses with SPAD-502 (Resende et al. unpublished data), suggesting a greater investment in light-harvesting structures under low-light conditions. This allocation pattern likely explains the higher foliar N levels observed under VLR despite lower total protein content, as more nitrogen was directed to chlorophyll and associated proteins rather than to bulk protein synthesis. Positive correlations between foliar N and chlorophyll content have also been reported in other species (Mizusaki et al. 2013 ; Yang et al. 2014a ; Schlichting et al. 2015 ). Proline levels exhibited divergent responses: they increased under HR in the control group but decreased under LR and HR in organic fertilizer plants. Proline accumulation differs with fertilization and light, suggesting that its role in osmoprotection or stress mitigation varies depending on nutrient status. Although proline stabilizes subcellular structures and acts as a redox buffer (Reddy et al. 2004 ; Molinari et al. 2007 ; Verslues and Sharma 2010 ), its primary role is to facilitate osmotic adjustment under water stress (Hong et al. 2000 ; Carvalho et al. 2013 ). SOD serves as the first enzymatic barrier against ROS, converting O 2 . - to H 2 O 2 (Alscher et al. 2002 ), which is then scavenged by CAT and various PODs to prevent the formation of highly reactive species, such as OH. (Kibinza et al. 2011 ; Shivashankara et al. 2016 ). PPO, abundant in plants, oxidizes phenolics into quinones and may act synergistically with POD (Krishna et al. 2008 ). Overall, SOD and PPO activities were significantly lower under HR, as were H 2 O 2 and superoxide levels. Such reductions in SOD and PPO under HR may reflect enzyme inhibition or a strategic shift in antioxidant defense, with other enzymes (POD, CAT) compensating depending on light intensity. CAT and POD showed complementary activity: CAT was more active under VLR and HR, while POD peaked under LR. Lipid peroxidation (TBARs index) was highest under HR. These patterns indicate that high radiation may trigger protein synthesis and lipid peroxidation as part of a protective response to light stress, while simultaneously reducing ROS accumulation through efficient scavenging mechanisms. This finding aligns with the observed increase in lipid peroxidation but lower ROS levels under HR, suggesting a dynamic regulation of defense pathways. These results also indicate increased ROS production in shaded environments (LR) and an effective response to oxidative stress under higher radiation. Interestingly, lower H 2 O 2 and superoxide levels under HR may reflect alternative antioxidant defenses, such as carotenoids, which deactivate singlet oxygen and other ROS. Indeed, S. jambos under HR exhibited significantly lower chlorophyll/carotenoid ratios (Resende et al. unpublished), highlighting the photoprotective role of these pigments in full sunlight conditions. Moreover, fertilization modulates these stress responses, likely by influencing nutrient-dependent metabolic pathways and antioxidant capacity. Contrasting results are reported in the literature regarding these enzymes activities. Yang et al. ( 2008 ) found increased SOD and CAT activities, and reduced POD activity in Picea asperata under high light conditions. The role of CAT in H 2 O 2 detoxification under intense radiation was demonstrated in Arabidopsis thaliana mutants, which exhibited elevated H 2 O 2 levels and cell death (Vandenabeele et al. 2004 ; Vanderauwera et al. 2005 ). Liu et al. ( 2016 ) observed significantly higher levels of hydrogen peroxide and superoxide radicals, along with increased activities of CAT, SOD, and APX, in Alnus species grown under full sunlight, with no significant changes in lipid peroxidation. In Torreya grandis , SOD activity was light-dependent, peaking at both the extremes of light levels, indicating that both excessive and insufficient light can impair plant metabolism (Tang et al. 2015 ). In the present study, gas exchange measurements were conducted under a standardized light intensity (1,000 µmol photons m -2 s -1 ) using the IRGA LED light chamber. The higher values of A , E, gs , and A /Ci observed in plants from shaded environments indicate an enhanced capacity for light utilization under limited irradiance. Furthermore, these results demonstrate the absence of photoinhibition, enabling shade-adapted plants to efficiently exploit the available light in environments where this resource is scarce. Shade plants in forest understories often experience brief, high-intensity sunflecks throughout the day, which contribute substantially to their daily carbon gain. These species are able to utilize sunflecks for positive net photosynthesis without suffering significant photoinhibition, due to rapid activation of energy dissipation mechanisms and other physiological adaptations (Way and Pearcy 2012 , Zhang et al. 2021 ). In contrast, elevated WUE in plants exposed to HR reflects efficient carbon assimilation relative to transpiration, indicating reduced water loss. Different patterns were reported for Piper aduncum , a pioneer species, which showed increased A and E and reduced WUE under higher light (Pacheco et al. 2013 ). Similarly, Alnus formosana , an invasive species, exhibited higher A , E , and g s under high radiation (Liu et al. 2016 ). Compared to the native species Alnus cremastogyne , A. formosana demonstrated superior photosynthetic and stomatal performance, highlighting functional advantages linked to invasiveness. In our study, high irradiance led to reduced net photosynthesis and increased stomatal limitation, reducing CO 2 assimilation and carboxylation efficiency. In contrast to our work, Silva et al. ( 2016 ) reported increased RuBisCo carboxylation efficiency ( A / C i ) under high light, suggesting species-specific responses to irradiance. These findings underscore the complexity of light-mediated physiological responses and the importance of ecological strategy in shaping gas exchange dynamics. Photosynthetic traits of plants from different successional stages were compared under contrasting light conditions (Zhang et al. 2015 ). Pioneer species showed greater reductions in photosynthetic rates under shade compared to mid- and late-successional species. This reflects the generally lower shade tolerance of early successional species and their reliance on open (under high light) environments for optimal carbon gain. Additional traits – such as lower light compensation points, reduced respiration rates, and higher apparent quantum yield – suggest that late-successional species are better adapted to low-light conditions. This response aligns with our findings, considering that light levels in the understory of mature forests, where these species grow and S. jambos is commonly found, can drop below 2% of full daylight (Chazdon and Fetcher 1984 ). Such environments favor species with enhanced light-use efficiency and physiological plasticity. The ΦPSII represents the fraction of absorbed light energy allocated to photochemical processes and typically shows higher values in plants exposed to low irradiance (Yang et al. 2025 ). Species adapted to high light intensities tend to exhibit lower effective quantum yield of PSII, as a larger portion of absorbed light is dissipated through non-photochemical pathways. This strategy helps to protect the photosynthetic apparatus from excess energy, minimizing photoinhibition (Yang et al. 2025 ). This pattern was confirmed in our study, where measured fluorescence parameters were generally higher in plants grown under VLR compared to those under HR. Previous studies support these findings. Zhang et al. ( 2015 ), for example, reported increased ΦPSII in response to reduced light intensity in five tree species; for Quercus aliena , shaded plants exhibited higher ΦPSII and ETR values (Xu et al. 2015 ). Enhanced photochemical efficiency under low light likely contributes to maintaining positive carbon balance in shaded environments, reinforcing the ecological advantage of high shade tolerance in S. jambos . Variations in gas exchange parameters across dehydration durations and light environments reveal a nuanced interaction between water availability and irradiance. Under HR, sharper declines in A , E , and g s after 20 days of dehydration reflect intensified physiological stress. Stomatal closure under drought is a primary adaptive response to minimize transpirational water loss. However, this response also restricts CO 2 uptake, thereby limiting photosynthetic carbon assimilation (Flexas et al. 2004 , 2006 , Oguz et al. 2022 ). Despite these reductions, the stability of WUE suggests a proportional decline in both CO 2 assimilation and water loss, reflecting balanced stomatal regulation. While stomatal closure explains the initial reductions in gas exchange, the observed drop in A / C i after 20 days of dehydration under HR points to additional non-stomatal limitations, likely linked to biochemical constraints on photosynthesis (Flexas and Medrano 2002 ). Moreover, the overall stability of A / C i suggests that such biochemical limitations only emerged under more severe stress, highlighting the species’ resilience to combined environmental pressures. Under high irradiance, the compounded effects of water deficit and excess energy intensify photoprotective demands, leading to increased non-photochemical energy dissipation and further constraining carbon assimilation (Nosalewicz et al. 2022 ). Our findings align with recent work by He et al. ( 2024 ), who reported similar patterns in tropical perennial trees: g s , A , and E d ropped significantly under drought, followed by strong recovery after rehydration. In our study, the substantial recovery of A , E , g s , and A / C i – especially under HR – along with the rapid restoration of gas exchange and water potential after rehydration, reveals the strong desiccation tolerance and hydraulic recovery mechanisms of S. jambos . The rebound in plant water potential, particularly under HR, further supports the species’ resilience to episodic drought, provided rehydration occurs. Such resilience and plasticity under fluctuating environmental conditions may help explain the invasive potential of S. jambos , especially in regions prone to intermittent drought and high irradiance. The results of our study allow us to conclude that S. jambos exhibits remarkable physiological and biochemical plasticity, enabling it to thrive under varying light intensities, fertilization regimes, and water availability. Light intensity strongly influenced nitrogen allocation, protein synthesis, antioxidant activity, and gas exchange. The species maintains photosynthetic efficiency and photochemical performance across contrasting light environments, deploys effective antioxidant and osmotic stress responses, and recovers rapidly from dehydration. Its flexible allocation of nitrogen and pigments, combined with tolerance to both shade and high irradiance, underpins its ecological versatility and likely contributes to its invasive potential. Understanding these adaptive mechanisms is essential for predicting their behavior under climate change and informing environmental management strategies. Declarations Acknowledgments The authors thank the Fundação de Amparo à Pesquisa do Estado de Minas Gerais (FAPEMIG) and the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for their financial support. Competing interests The authors have no relevant financial or non-financial interests to disclose. Author Contributions Both authors contributed to the conception and design of the study. Material preparation, data collection, and analysis were performed by Cristiano Ferrara de Resende. The first draft of the manuscript was written by Cristiano Ferrara de Resende and both authors commented on previous versions. 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1","display":"","copyAsset":false,"role":"figure","size":148941,"visible":true,"origin":"","legend":"\u003cp\u003eGas exchange parameters in \u003cem\u003eSyzygium jambos\u003c/em\u003e plants after 11 months of growth under three light environments (very low radiation – VLR, low radiation – LR, and high radiation – HR), subjected to two fertilization systems (control group and bovine manure-fertilized). \u003cstrong\u003ea)\u003c/strong\u003e CO\u003csub\u003e2\u003c/sub\u003e assimilation rate (\u003cem\u003eA\u003c/em\u003e, µmol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e); \u003cstrong\u003eb)\u003c/strong\u003e transpiration (\u003cem\u003eE\u003c/em\u003e, mmol m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e); \u003cstrong\u003ec)\u003c/strong\u003e stomatal conductance (\u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e, mol H\u003csub\u003e2\u003c/sub\u003eO m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e); \u003cstrong\u003ed)\u003c/strong\u003e substomatal CO\u003csub\u003e2\u003c/sub\u003e concentration (\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e, µmol mol\u003csup\u003e-1\u003c/sup\u003e); \u003cstrong\u003ee)\u003c/strong\u003e water use efficiency (WUE, µmol CO\u003csub\u003e2\u003c/sub\u003e mmol\u003csup\u003e-1\u003c/sup\u003e H\u003csub\u003e2\u003c/sub\u003eO); \u003cstrong\u003ef)\u003c/strong\u003e intrinsic water use efficiency (WUE\u003csub\u003eint\u003c/sub\u003e, µmol CO\u003csub\u003e2 \u003c/sub\u003emol\u003csup\u003e-1\u003c/sup\u003e H\u003csub\u003e2\u003c/sub\u003eO); \u003cstrong\u003eg)\u003c/strong\u003e carboxylation efficiency (\u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e, µmol m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e Pa\u003csup\u003e-1\u003c/sup\u003e). Uppercase letters compare the effect of light intensity within each fertilization system (control and bovine manure-fertilized), while lowercase letters compare the effect of fertilization within each light intensity. Means followed by the same letters do not differ from each other according to the Scott-Knott test at a 5% probability\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8196016/v1/16192f7301c738552e6d3c0f.png"},{"id":97140188,"identity":"95e42a0c-c81d-43c2-8451-692d398cbdd2","added_by":"auto","created_at":"2025-12-01 10:04:05","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":85212,"visible":true,"origin":"","legend":"\u003cp\u003eFluorescence parameters in \u003cem\u003eS. jambos\u003c/em\u003e plants after 11 months of development under three light environments (very low radiation – VLR, low radiation – LR, and high radiation – HR), subjected to two fertilization systems (control group and bovine manure-fertilized). \u003cstrong\u003ea)\u003c/strong\u003e Energy capture efficiency of excitation by the open reaction centers of PSII (Fv'/Fm'); \u003cstrong\u003eb)\u003c/strong\u003e effective quantum yield of PSII electron transport (ΦPSII); \u003cstrong\u003ec)\u003c/strong\u003e photochemical quenching coefficient (q\u003csub\u003eP\u003c/sub\u003e); \u003cstrong\u003ed)\u003c/strong\u003e apparent electron transport rate (ETR). Uppercase letters compare the effect of light intensity within each fertilization system (control and bovine manure-fertilized), and lowercase letters compare the effect of fertilization within each light intensity. Means followed by the same letters do not differ statistically according to the Scott-Knott test at a 5% probability\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8196016/v1/17711e7074300250c9de6932.png"},{"id":97027947,"identity":"2d85c11e-aef4-4587-86cf-7369cf196c74","added_by":"auto","created_at":"2025-11-29 00:39:06","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":288229,"visible":true,"origin":"","legend":"\u003cp\u003eVisual appearance of \u003cem\u003eSyzigium jambos\u003c/em\u003e plants subjected to 10 and 20 days of dehydration under very low radiation (VLR) and high radiation (HR). Progressive leaf wilting and stem curvature are more pronounced under VLR after 20 days, indicating greater physiological stress compared to HR conditions\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-8196016/v1/462fa86af0687470aa2bb6f4.png"},{"id":100360183,"identity":"37dfb825-49a4-40cc-ab74-bfa920ae86a4","added_by":"auto","created_at":"2026-01-16 07:37:52","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2231113,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8196016/v1/dd4dd0b9-e2ef-4440-9276-cb45cad7b5d1.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Physiological and biochemical plasticity as a driver of invasiveness in Syzygium jambos (L.) Alston","fulltext":[{"header":"Introduction","content":"\u003cp\u003eInvasive plant species are among the most transformative agents in tropical ecosystems, often reshaping community structure and ecosystem function. Their ecological success is frequently attributed to their ability to tolerate and thrive under a wide range of environmental conditions (Moura et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Hern\u0026aacute;ndez-Fern\u0026aacute;ndez et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). \u003cem\u003eSyzygium jambos\u003c/em\u003e (L.) Alston (Myrtaceae), commonly known as rose apple, exemplifies this adaptability. Native to Tropical Asia, this fast-growing tree species can exceed 12 meters in height and is widely recognized for its capacity to colonize both disturbed and undisturbed tropical habitats. Its rapid establishment in open areas and mature forests has led to significant alterations in native plant assemblages, particularly through the formation of dense canopies that suppress light-dependent species (Horvitz et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Aide et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Lugo \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Brown et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Cramer et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Morales \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn tropical forests, where light is a limiting and heterogeneously distributed resource, the ability to optimize light use is crucial for plant performance. Light availability drives key physiological processes, including photosynthesis, antioxidant metabolism, and nutrient uptake (Valladares et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; dos Anjos et al. 2015; Liu et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The interplay between light intensity and soil nutrient availability can significantly shape competitive interactions among species, particularly in environments where resource partitioning is crucial for growth and survival (Ferreira et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). While native species have evolved strategies to cope with such variability, invasive plants often exhibit enhanced phenotypic plasticity, enabling them to outperform native flora under fluctuating conditions (Davidson et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eInvasive plants also tend to exhibit physiological and biochemical traits that enhance their resilience under environmental stress, contributing to their competitive success. Traits such as increased photosynthetic efficiency, flexible stomatal regulation, and robust antioxidant systems enable these species to maintain functional performance across diverse light, nutrient, and water conditions (McAlpine et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pint\u0026oacute;-Marijuan and Munn\u0026eacute;-Bosch \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Sanders et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Chlorophyll fluorescence is a sensitive tool for assessing photochemical efficiency and has been widely used to assess stress responses in invasive taxa (Zhou et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Additionally, the activity of antioxidant enzymes \u0026ndash; including superoxide dismutase (SOD), catalase (CAT), peroxidase (POD), and polyphenol oxidase (PPO) \u0026ndash; plays a central role in mitigating oxidative damage caused by reactive oxygen species (ROS), which tend to accumulate under abiotic stress such as drought, nutrient limitation, and excess light (Wang et al. 2023). These physiological mechanisms, when coupled with phenotypic plasticity, may explain the ability of \u003cem\u003eS. jambos\u003c/em\u003e to persist and dominate in diverse tropical environments.\u003c/p\u003e\u003cp\u003eBeyond ecological disruption, biological invasions impose substantial economic costs. A global synthesis estimated that the cumulative cost of invasive species reached at least USD 1.28 trillion between 1970 and 2017, with annual costs tripling every decade and peaking at USD 162.7\u0026nbsp;billion in 2017 alone (Diagne et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Notably, damage-related expenses \u0026ndash; such as loss of ecosystem services and agricultural productivity \u0026ndash; far exceed management investments, particularly for plant taxa, which remain underrepresented in economic assessments. In Brazil, the economic burden of biological invasions has recently been quantified for the first time, revealing a minimum cost of USD 105.53\u0026nbsp;billion over 35 years, with an overwhelming 99% of this amount attributed to damages and losses rather than management efforts (Adelino et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Given that only 16 invasive species were included in this estimate \u0026ndash; out of at least 460 recognized invaders \u0026ndash; the actual economic impact is likely much higher. An updated national inventory has since identified 444 invasive non-native species in Brazil, including 188 plants, with trees being the most frequent life form among them (Zenni et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This reinforces the relevance of investigating arboreal invaders such as \u003cem\u003eS. jambos\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eThese findings underscore the urgency of understanding the physiological mechanisms that underpin invasion success, especially in species like \u003cem\u003eS. jambos\u003c/em\u003e, whose ecological dominance is matched by its potential economic impact. This study aimed to investigate the physiological and biochemical traits of \u003cem\u003eS. jambos\u003c/em\u003e under varying environmental conditions, in order to identify the features most closely associated with its invasive behavior. By linking ecophysiological performance to invasion potential, we seek to contribute to a more predictive framework for managing exotic species in tropical forest ecosystems.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003ePlant material and experimental design\u003c/h2\u003e\u003cp\u003eFruits of \u003cem\u003eS. jambos\u003c/em\u003e were collected from two specimens at the Institute of Biological Sciences (ICB) of the Federal University of Juiz de Fora (UFJF), Minas Gerais, Brazil. Seeds were extracted, placed in Petri dishes with filter paper, and moistened with 20 mL of distilled water to induce imbibition and germination. Germinated seeds were transplanted into 200 mL plastic cups containing Plantmax Hortali\u0026ccedil;as HT\u0026reg; substrate, kept in shade, and irrigated twice weekly. All seeds developed into seedlings, which were standardized by height (10\u0026ndash;12 cm) and leaf count (8\u0026ndash;10 leaves) prior to experimental setup.\u003c/p\u003e\u003cp\u003eSeedlings were transferred to 25 L pots filled with one of three substrate treatments: 1) control \u0026ndash; soil and sand (3:2, v/v); 2) bovine manure \u0026ndash; soil, sand, and aged bovine manure (3:2:1, v/v/v); and 3) chemical fertilizer \u0026ndash; soil and sand (3:2, v/v)\u0026thinsp;+\u0026thinsp;50 g of Forth Frutas\u0026reg; (12% N, 5% P, 15% K). The pots were placed in a greenhouse under shade conditions (4% of ambient light, ~\u0026thinsp;100 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e of photosynthetically active radiation \u0026ndash; PAR), achieved using Sombrite\u0026reg; screens of different mesh densities. Full sunlight (~\u0026thinsp;2,400 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e of PAR on clear days) was used as the reference for 100% irradiance. Plants remained in this environment for ~\u0026thinsp;8 months to allow growth and tissue development. All plants in the chemically fertilized treatment died during this period, leaving only the control group and the bovine manure groups.\u003c/p\u003e\u003cp\u003eFor the new experimental setup, plants from these two groups were standardized by height (30\u0026ndash;50 cm, mean of 40 cm) and the number of leaves (12\u0026ndash;18 leaves, mean of 16). They were then assigned to three light treatments: very low radiation (VLR) \u0026ndash; 4% of ambient light (~\u0026thinsp;100 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e PAR); low radiation (LR) \u0026ndash; 9% of ambient light (~\u0026thinsp;220 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e PAR); and high radiation (HR) \u0026ndash; full sunlight (~\u0026thinsp;2,400 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e PAR). The experiments were conducted using a completely randomized factorial design (2 fertilization types \u0026times; 3 light levels), totaling six treatments. All plants were irrigated at least twice a week and maintained under these conditions for 11 months, after which leaf samples were collected for biochemical analysis.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003ePhysicochemical properties and fertility assessment of substrates\u003c/h3\u003e\n\u003cp\u003ePrior to the experiment, random samples of the three substrates were collected to assess particle size distribution and nutrient composition. After 11 months of plant growth, three samples from each of the six final treatments were collected and analyzed for their nutritional and chemical attributes. Nutrient extraction followed standard protocols: Mehlich 1 (0.05 M HCl\u0026thinsp;+\u0026thinsp;0.0125 M H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e) for K, P, Zn, Fe, Mn, and Cu; 1M KCl for Ca, Mg, and Al; hot water for B; monocalcium phosphate in acetic acid for S; pH in SMP buffer solution for potential acidity (H\u0026thinsp;+\u0026thinsp;Al). Additional parameters included: pH in water (1:2.5), sum of exchangeable bases (SB), cation exchange capacity at pH 7.0 (T), effective cation exchange capacity (t), base saturation index (V), aluminum saturation index (m), organic matter (OM; via oxidation with 4N Na\u003csub\u003e2\u003c/sub\u003eCr\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e7\u003c/sub\u003e\u0026thinsp;+\u0026thinsp;10N H\u003csub\u003e2\u003c/sub\u003eSO\u003csub\u003e4\u003c/sub\u003e), and remaining phosphorus (P-Rem).\u003c/p\u003e\n\u003ch3\u003eBiochemical and enzymatic analysis\u003c/h3\u003e\n\u003cp\u003eCrude extracts for total protein and enzymatic activity determinations \u0026ndash; superoxide dismutase (SOD; EC 1.15.1.1), catalase (CAT; EC 1.11.1.6), peroxidase (POD; EC 1.11.1.7), and polyphenol oxidase (PPO; EC 1.10.3.1, EC 1.10.3.2, and EC 1.14.18.1) \u0026ndash; were obtained by macerating four leaf discs (0.8 cm\u003csup\u003e2\u003c/sup\u003e; 0.07\u0026ndash;0.1 g each) in liquid nitrogen, followed by homogenization in 5 mL of buffer containing 0.1 M potassium phosphate (pH 6.8), 0.1 mM EDTA, and 1 mM PMSF (Peixoto et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). The homogenate was filtered through gauze and centrifuged (10,000 \u003cem\u003eg\u003c/em\u003e, 15 min, 4\u0026deg;C). Soluble protein content was determined using the method of Lowry et al. (\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1951\u003c/span\u003e), with the Folin-Ciocalteu reagent. The protein concentration in each replicate was used to calculate the specific activity of the enzymes. SOD activity was measured according to Del Longo et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1993\u003c/span\u003e), with reactions conducted at 25\u0026deg;C under illumination by a 15W fluorescent lamp (Giannopolitis and Ries \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1977\u003c/span\u003e). Photoreduction of nitro blue tetrazolium (NBT) was monitored at 560 nm, and one unit of activity was defined as the amount of enzyme required to inhibit 50% of NBT reduction (Beauchamp and Fridovich \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1971\u003c/span\u003e). CAT activity was measured by monitoring H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e decomposition at 240 nm (Havir and McHale \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1987\u003c/span\u003e), using a molar extinction coefficient of 36 M\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (Anderson et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). POD and PPO activities were estimated according to Kar and Mishra (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1976\u003c/span\u003e), with absorbance measured at 420 nm. A molar extinction coefficient of 2.47 mM\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e was used for calculations (Chance and Maehley 1954). Proline quantification was performed according to Bates et al. (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1973\u003c/span\u003e). Six leaf discs (0.8 cm\u003csup\u003e2\u003c/sup\u003e; 0.1\u0026ndash;0.16 g each) were macerated in liquid nitrogen and homogenized in 10 mL of 3% (w/v) sulfosalicylic acid, then filtered through Whatman No. 2 paper. For each sample, 2 mL of filtrate was combined with 2 mL of acidic ninhydrin and 2 mL of acetic acid, then incubated at 100\u0026deg;C for 1 hour. After cooling in an ice bath, the chromophore was extracted with toluene and separated by decantation. Absorbance was measured at 520 nm, and proline concentration was determined using a standard curve. Hydrogen peroxide (H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e) content was determined using a modified ferrous ammonium sulfate/xylenol orange (FOX) assay, with absorbance measured at 560 nm (Gay and Gebicki \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Approximately 0.2 g of leaf tissue (7\u0026ndash;11 disks, 0.8 cm\u003csup\u003e2\u003c/sup\u003e each) was used, and H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e concentration was calculated from a standard curve prepared with authentic H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e solutions. Superoxide anion (O\u003csub\u003e2\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e.) content was measured following Mohammadi and Karr (\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), with modifications. Leaf samples (0.3 g; 40 discs, 0.2 cm\u0026sup2; each) were incubated in reaction medium, and superoxide production was quantified based on adrenochrome formation (Misra and Fridovich \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1971\u003c/span\u003e), using a molar extinction coefficient of 4.0 x 10\u003csup\u003e3\u003c/sup\u003e M\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (Boveris \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). Lipid peroxidation was assessed by quantifying malondialdehyde (MDA) produced after reaction with thiobarbituric acid (TBA) (Cakmak and Horst \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). Approximately 0.2 g of leaf tissue (25\u0026ndash;38 disks, 0.2 cm\u003csup\u003e2\u003c/sup\u003e each) was used per sample. Absorbance was recorded at 532 and 600 nm. MDA-TBA concentration was calculated using a molar extinction coefficient of 155 mM\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (Heath and Packer \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1968\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eGas exchange and chlorophyll fluorescence\u003c/h3\u003e\n\u003cp\u003eGas exchange and chlorophyll \u003cem\u003ea\u003c/em\u003e fluorescence measurements were conducted after 11 months of plant exposure to different light environments. Analyses were performed between 08:00 and 12:00 h using an infrared gas analyzer (LI-6400XT, Li-Cor, Lincoln, NE, USA) with an LED fluorescence chamber (6400-02B, Li-Cor). Fully expanded, healthy leaves from the fourth or fifth node were selected for use. The leaf chamber was set to a photon flux density of 1,000 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e and ambient CO\u003csub\u003e2\u003c/sub\u003e concentration (~\u0026thinsp;400 \u0026micro;mol mol\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Leaf temperatures averaged 29\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C under HR and 34\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u0026deg;C under VLR and LR. Measured parameters included net photosynthetic rate (\u003cem\u003eA\u003c/em\u003e), transpiration rate (\u003cem\u003eE\u003c/em\u003e), stomatal conductance (\u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e), and intercellular CO\u003csub\u003e2\u003c/sub\u003e concentration (\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e). Derived indices estimated were the ratio of intercellular to external CO\u003csub\u003e2\u003c/sub\u003e concentration (\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ea\u003c/sub\u003e), water-use efficiency (WUE\u0026thinsp;=\u0026thinsp;\u003cem\u003eA\u003c/em\u003e/\u003cem\u003eE\u003c/em\u003e) (Ou et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), intrinsic WUE (WUE\u003csub\u003eint\u003c/sub\u003e = \u003cem\u003eA\u003c/em\u003e/\u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e), and carboxylation efficiency (\u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e). Fluorescence was assessed on light-adapted leaves exposed to a saturating pulse (6,000 \u0026micro;mol photons m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, 0.8 s) to determine steady-state fluorescence (F\u003csub\u003es\u003c/sub\u003e) and maximum fluorescence under light (F\u003csub\u003em\u003c/sub\u003e\u0026rsquo;). Far-red light was then applied to measure the minimum fluorescence of light-adapted leaves (F\u003csub\u003eo\u003c/sub\u003e\u0026rsquo;). Based on Genty et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1989\u003c/span\u003e), the following parameters were calculated: effective quantum yield of PSII electron transport (Φ\u003csub\u003ePSII\u003c/sub\u003e = (F\u003csub\u003em\u003c/sub\u003e\u0026rsquo; \u0026ndash; F\u003csub\u003es\u003c/sub\u003e)/F\u003csub\u003em\u003c/sub\u003e\u0026rsquo;), efficiency of excitation energy capture by open PSII reaction centers (F\u003csub\u003ev\u003c/sub\u003e\u0026rsquo;/F\u003csub\u003em\u003c/sub\u003e\u0026rsquo; = (F\u003csub\u003em\u003c/sub\u003e\u0026rsquo; \u0026ndash; F\u003csub\u003eo\u003c/sub\u003e\u0026rsquo;)/F\u003csub\u003em\u003c/sub\u003e\u0026rsquo;), the photochemical quenching (q\u003csub\u003eP\u003c/sub\u003e = (F\u003csub\u003em\u003c/sub\u003e\u0026rsquo; \u0026ndash; F\u003csub\u003es\u003c/sub\u003e)/(F\u003csub\u003em\u003c/sub\u003e\u0026rsquo; \u0026ndash; F\u003csub\u003eo\u003c/sub\u003e\u0026rsquo;)), and the apparent electron transport rate (ETR\u0026thinsp;=\u0026thinsp;Φ\u003csub\u003ePSII\u003c/sub\u003e\u0026thinsp;\u0026times;\u0026thinsp;PAR \u0026times; 0.5 \u0026times; 0.84, where PAR is the photosynthetically active radiation incident on the leaf, 0.5 represents the fraction of photons allocated to photosystem I and photosystem II, and 0.84 is the average fraction of incident light absorbed by the leaf).\u003c/p\u003e\n\u003ch3\u003eLeaf nutrient contents\u003c/h3\u003e\n\u003cp\u003eMacro- and micronutrient contents in leaves were determined at the end of the experiments. Samples were dried at 65\u0026deg;C to constant weight, ground, and analyzed using standardizing methodologies. The nutrients evaluated included N, P, K, Ca, Mg, S, B, Cu, Mn, Zn, and Fe. Nitrogen (N) content was determined using the sulfuric digestion method followed by quantification via the Kjeldahl procedure. For K, S, Fe, Ca, Zn, Mg, Cu, and Mn, tissue samples were digested in a nitric-perchloric solution (3:1; v:v). Boron (B) was extracted using a hydrochloric acid (HCl) solution. Nutrient concentrations were determined by atomic absorption spectrophotometry (for K, Ca, Mg, Fe, Cu, Mn, and Zn) and colorimetric spectrophotometry (for P, S, and B).\u003c/p\u003e\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003eDrought stress experiment\u003c/h2\u003e\u003cp\u003eAfter 11 months of growth, plants from the control group under the two most contrasting light conditions \u0026ndash; VLR and HR \u0026ndash; were selected for a drought stress experiment. These plants were subjected to a progressive dehydration protocol, in which irrigation was withheld for 10 or 20 days, while a well-watered control group continued to receive daily irrigation. This design resulted in six distinct treatments. Following the drought phase, all plants were rehydrated and maintained under optimal watering conditions.\u003c/p\u003e\u003cp\u003eLeaf samples were collected at the end of each drought period (10 or 20 days without irrigation) and one week after rehydration for physiological assessments. These included measurements of gas exchange parameters and antioxidant enzyme activities, as previously described. Soil moisture and temperature were continuously monitored throughout the experiment. Predawn leaf water potential was determined during both dehydration and rehydration phases using a Scholander-type pressure chamber, as described by Turner (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1988\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003eStatistical analysis\u003c/h2\u003e\u003cp\u003eData were first tested for homogeneity of variances (Cochran-Bartlett test) and normality (Lilliefors). Once these assumptions were met, data were subjected to analysis of variance (ANOVA), and means were grouped using the Scott-Knott test at a 5% probability level, performed with SAEG software (version 9.1).\u003c/p\u003e\u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003eChemical analyses of substrates\u003c/h2\u003e\u003cp\u003eBefore the cultivation, the substrate pH was near neutral in the control group (6.9) and slightly acidic in the bovine manure-fertilized group (6.2). In contrast, the chemically fertilized substrate was much more acidic (pH 4.5) (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Potential acidity (H\u0026thinsp;+\u0026thinsp;Al) was also markedly higher in the chemically fertilized material \u0026ndash; 4.5 times greater than the control and 3.6 times greater than the manure-fertilized group. Macronutrient (K, P, Ca, Mg, S) and micronutrient (Mn, B) levels were highest in the chemically fertilized substrate, followed by the manure-fertilized and control groups. Fe and Cu contents decreased in the order: chemically fertilized, control, and manure fertilized. Zn and organic matter (OM) were most abundant in the manure-fertilized substrate. Despite differences in P levels, P-Rem and base saturation (V) were similar across treatments. Confirmed by the aluminum saturation index (m), Al was detected only in the chemically fertilized substrate. Although classified as different soil types \u0026ndash; medium texture (control and chemical) and clayey texture (manure) \u0026ndash; the proportions of clay, silt, and sand were similar among them, with sand predominating.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eChemical characterization and granulometry of substrates used in the experiment. Parameters: pH of the substrates (pH); contents of potassium (K), phosphorus (P), calcium (Ca), magnesium (Mg), aluminum (Al), zinc (Zn), iron (Fe), manganese (Mn), copper (Cu), boron (B), sulfur (S), remaining phosphorus (P-Rem), organic matter (OM); potential acidity (H\u0026thinsp;+\u0026thinsp;Al); sum of exchangeable bases (SB), effective cation exchange capacity (CEC) (t), and CEC at pH 7.0 (T); base saturation index (V) and aluminum saturation index (m); clay, silt, and sand. Soil type 2 \u0026ndash; medium texture; soil type 3 \u0026ndash; clayey texture\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eParameter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e\u003cp\u003eSubstrate\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eControl\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eChemical fertilization\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eBovine manure\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003epH\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eK (\u003c/b\u003emg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e30.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e720.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e396.00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eP (\u003c/b\u003emg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e10.24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e121.49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e18.58\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCa\u003c/b\u003e (cmol dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e12.71\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5.30\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMg\u003c/b\u003e (cmol dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.58\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.20\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eAl\u003c/b\u003e (cmol dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eZn\u003c/b\u003e (mg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e26.41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e58.29\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e102.24\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eFe\u003c/b\u003e (mg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e82.67\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e96.36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e66.68\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMn\u003c/b\u003e (mg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e33.22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e67.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e59.86\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCu\u003c/b\u003e (mg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.77\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.96\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.68\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eB\u003c/b\u003e (mg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.84\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.50\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eS\u003c/b\u003e (mg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e37.91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e135.64\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e91.28\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eP-Rem\u003c/b\u003e (mg L\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e15.94\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e12.93\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e14.17\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eO.M\u003c/b\u003e (dag kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.87\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.61\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.84\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eH\u0026thinsp;+\u0026thinsp;Al\u003c/b\u003e (cmolc dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e7.04\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.94\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSB\u003c/b\u003e (cmolc dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e19.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.52\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003et\u003c/b\u003e (cmolc dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e19.34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.52\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eT\u003c/b\u003e (cmolc dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6.73\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e26.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10.46\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eV (%)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e76.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e73.09\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e81.41\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003em (%)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.03\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.00\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eClay\u003c/b\u003e (dag kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e37\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSilt\u003c/b\u003e (dag kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSand\u003c/b\u003e (dag kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e69\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e61\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e59\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eClassification\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType 2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eType 2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eType 3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eSince plants treated with chemical fertilizer died before the start of light intensity treatments, only the control and bovine manure-fertilized substrates were evaluated. Performed 11 months after plant development, nutritional analyses of the substrates showed that in the control group, radiation intensity had no significant effect on most soil parameters, except for S levels, which decreased with increasing light intensity (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In contrast, bovine manure-fertilized substrates showed marked differences in the concentration of essential elements across light environments. After nearly one year of cultivation, substrate pH under HR was significantly higher (mean 7.03) than in LR and VLR. Under VLR, levels of K, P, Mg, B, sum of exchangeable bases (SB), and cation exchange capacity (CEC) at pH 7.0 (T) were significantly more elevated, with no differences between LR and HR. Ca, Zn, Fe, Mn, remaining phosphorus (P-Rem), organic matter (OM), and base saturation index (V) showed no significant variation across light treatments. Cu was lower under HR, and potential acidity (H\u0026thinsp;+\u0026thinsp;Al) was reduced under VLR. S levels mirrored those in the control, decreasing with higher radiation. Al was not detected in any of the samples. No significant differences in pH or in the Zn, Fe, Cu, and S concentrations were found between control and bovine manure-fertilized substrates across light environments. After 11 months, substrates prepared with manure generally showed higher levels of K, P, Ca, Mg, P-Rem, OM, SB, T, and V. Compared to the control, higher concentrations of Mn and B were detected only in the fertilized material under VLR. In VLR and LR conditions, H\u0026thinsp;+\u0026thinsp;Al levels were higher in control than in those with organic amendment.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eChemical attributes of substrates of \u003cem\u003eSyzygium jambos\u003c/em\u003e plants after 11 months of development in three light environments (very low radiation \u0026ndash; VLR, low radiation \u0026ndash; LR, and high radiation \u0026ndash; HR), subjected to two fertilization systems (control group and bovine manure-fertilized). Parameters: pH of the substrates (pH); contents of potassium (K), phosphorus (P), calcium (Ca), magnesium (Mg), zinc (Zn), iron (Fe), manganese (Mn), copper (Cu), boron (B), sulfur (S), remaining phosphorus (P-Rem), organic matter (OM); potential acidity (H\u0026thinsp;+\u0026thinsp;Al); sum of exchangeable bases (SB), cation exchange capacity (CEC) at pH 7.0 (T); and base saturation index (V). Uppercase letters compare the effect of radiation within each fertilization system, while lowercase letters compare the effect of fertilization within each light intensity. Data are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eParameter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e\u003cp\u003eControl\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003eBovine manure\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003epH\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6.87\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.30 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e6.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e7.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eK (\u003c/b\u003emg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e26.67\u0026thinsp;\u0026plusmn;\u0026thinsp;1.76 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e26.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e25.33\u0026thinsp;\u0026plusmn;\u0026thinsp;2.40 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e199.33\u0026thinsp;\u0026plusmn;\u0026thinsp;37.81 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e110.00\u0026thinsp;\u0026plusmn;\u0026thinsp;12.49 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e64.67\u0026thinsp;\u0026plusmn;\u0026thinsp;6.77 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eP (\u003c/b\u003emg dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e10.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e9.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.66 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.44\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e21.37\u0026thinsp;\u0026plusmn;\u0026thinsp;2.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e16.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" 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dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.89\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.68\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSB\u003c/b\u003e (cmolc dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e5.30\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7.44\u0026thinsp;\u0026plusmn;\u0026thinsp;0.63 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e6.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e5.96\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eT\u003c/b\u003e (cmolc dm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e7.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6.89\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e9.14\u0026thinsp;\u0026plusmn;\u0026thinsp;0.59 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e8.18\u0026thinsp;\u0026plusmn;\u0026thinsp;0.26 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e7.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eV\u003c/b\u003e (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e74.39\u0026thinsp;\u0026plusmn;\u0026thinsp;1.24 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e72.51\u0026thinsp;\u0026plusmn;\u0026thinsp;1.24 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e75.14\u0026thinsp;\u0026plusmn;\u0026thinsp;1.05 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e81.40\u0026thinsp;\u0026plusmn;\u0026thinsp;3.29 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e80.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e77.99\u0026thinsp;\u0026plusmn;\u0026thinsp;1.80 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eComparisons between initial and post-cultivation substrate fertility revealed notable shifts (Tables\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e and \u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). pH decreased in unfertilized substrates but increased in those with bovine manure, mirroring trends in potential acidity (H\u0026thinsp;+\u0026thinsp;Al). Macronutrients (K, P, Ca, Mg, S) and parameters SB, t, and T declined more sharply in manure-amended substrates, proportionally to light intensity. Micronutrient levels (Zn, Fe, Mn, Cu, B) fluctuated without consistent patterns. After 11 months, OM increased in control but declined in substrates with manure addition, while P-Rem showed the opposite trend.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003eFoliar nutrient concentrations\u003c/h2\u003e\u003cp\u003eThe results for foliar nutrient concentrations are summarized in Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e. In the control group, the concentrations of P, K, Ca, Mg, B, Mn, and Fe did not vary significantly with light intensity. However, foliar N decreased as light radiation increased, with significant differences observed in both control and manure-fertilized plants. In control plants, S was highest under VLR, while Cu and Zn peaked under HR. In manure-fertilized plants, most nutrients varied significantly across light treatments, except for Fe, which remained stable. HR conditions favored the accumulation of P, Mg, B, Cu, and Zn, while the opposite was observed for K, Ca, and S (higher concentrations in VLR). Across all light environments, S, Zn, and Fe showed no significant differences between control and fertilized groups. In general, fertilized plants showed higher levels of N, P, and K, although these differences were not always statistically significant. Notably, B and Cu were significantly higher in fertilized plants under HR. Control plants had higher Ca and Mg overall, except for Mg under HR.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eFoliar nutrient concentrations in \u003cem\u003eSyzygium jambos\u003c/em\u003e plants after 11 months of growth under three light environments (very low radiation \u0026ndash; VLR; low radiation \u0026ndash; LR; and high radiation \u0026ndash; HR) and two fertilization systems (control group and bovine manure-fertilized). Uppercase letters compare the effect of radiation within each fertilization system, while lowercase letters compare the effect of fertilization within each light intensity. Data are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eParameter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e\u003cp\u003eControl\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003eBovine manure\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eN\u003c/b\u003e (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.78\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Cb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Ca\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eP\u003c/b\u003e (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eK\u003c/b\u003e (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 Ca\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCa\u003c/b\u003e (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.88\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.84\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.77\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.57\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Bb\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMg\u003c/b\u003e (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eS\u003c/b\u003e (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.14\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eB\u003c/b\u003e (ppm)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e15.3\u0026thinsp;\u0026plusmn;\u0026thinsp;2.5 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e11.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e24.3\u0026thinsp;\u0026plusmn;\u0026thinsp;6.8 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e19.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e22.3\u0026thinsp;\u0026plusmn;\u0026thinsp;4.5 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e35.8\u0026thinsp;\u0026plusmn;\u0026thinsp;3.0 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCu\u003c/b\u003e (ppm)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.53\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.87\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMn\u003c/b\u003e (ppm)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e48.2\u0026thinsp;\u0026plusmn;\u0026thinsp;35.2 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e14.8\u0026thinsp;\u0026plusmn;\u0026thinsp;3.1 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.1\u0026thinsp;\u0026plusmn;\u0026thinsp;2.8 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10.7\u0026thinsp;\u0026plusmn;\u0026thinsp;3.5 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e167.9\u0026thinsp;\u0026plusmn;\u0026thinsp;76.6 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e21.5\u0026thinsp;\u0026plusmn;\u0026thinsp;9.5 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eZn\u003c/b\u003e (ppm)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e13.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e13.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e20.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e12.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e11.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e21.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eFe\u003c/b\u003e (ppm)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e131.3\u0026thinsp;\u0026plusmn;\u0026thinsp;19.3 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e131.3\u0026thinsp;\u0026plusmn;\u0026thinsp;38.5 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e88.4\u0026thinsp;\u0026plusmn;\u0026thinsp;4.0 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e129.7\u0026thinsp;\u0026plusmn;\u0026thinsp;5.5 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e110.2\u0026thinsp;\u0026plusmn;\u0026thinsp;6.2 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e93.6\u0026thinsp;\u0026plusmn;\u0026thinsp;5.2 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003eBiochemical analyses\u003c/h2\u003e\u003cp\u003eRegardless of the fertilization method or measurement unit, plants under HR had significantly higher total protein content (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). In the control group, proline accumulated more under HR, while in fertilized plants, it was higher under VLR. SOD and PPO activities were lowest under HR, especially in fertilized plants, which showed significant variation across light treatments. CAT activity remained unchanged in fertilized plants but decreased under LR in the control group, whereas POD activity peaked under LR in both groups. Overall, hydrogen peroxide (H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e) and superoxide (O\u003csub\u003e2\u003c/sub\u003e.\u003csup\u003e\u0026minus;\u003c/sup\u003e) levels were lowest under HR, while MDA content was highest under HR, regardless of fertilization. Comparing control and fertilized plants, protein content differed significantly only under HR, with higher values in the control group. Proline was higher in fertilized plants under VLR and in control plants under HR. SOD activity differed only under LR, being lower in fertilized plants. CAT activity varied under VLR and HR, with fertilized plants showing the lowest values. POD and PPO activities differed only under VLR, with higher POD in control and higher PPO in fertilized plants. Hydrogen peroxide, superoxide, and MDA levels were generally similar between groups, except for MDA under HR, which was higher in fertilized plants.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eBiochemical attributes of \u003cem\u003eSyzygium jambos\u003c/em\u003e plants after 11 months of growth under three light environments (very low radiation \u0026ndash; VLR, low radiation \u0026ndash; LR, and high radiation \u0026ndash; HR) and two fertilization systems (control group and bovine manure-fertilized). Total protein content is presented on a fresh mass basis (mg g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e FM) and area basis (mg cm\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e); proline content (\u0026micro;mol g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e FM); enzymatic activities of SOD (units mg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e protein), CAT (\u0026micro;mol of H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e min\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e mg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e protein), POD (\u0026micro;mol mg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e protein), and PPO (\u0026micro;mol mg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e protein); and contents of peroxide (nmol g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e FM), superoxide (nmol g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e FM), and malondialdehyde (MDA; \u0026micro;mol g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e FM). Uppercase letters compare the effect of radiation within each fertilization system, while lowercase letters compare the effect of fertilization within each light intensity. Data are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eParameter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e\u003cp\u003eControl\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c7\" namest=\"c5\"\u003e\u003cp\u003eBovine manure\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eProteins (/mass)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e26.0\u0026thinsp;\u0026plusmn;\u0026thinsp;1.8 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e23.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.4 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e93.6\u0026thinsp;\u0026plusmn;\u0026thinsp;2.5 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e20.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e20.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e81.8\u0026thinsp;\u0026plusmn;\u0026thinsp;3.5 Ab\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eProteins (/area)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.53\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e2.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16 Ab\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eProline\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.34\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.43\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Bb\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSOD\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e24.47\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e22.40\u0026thinsp;\u0026plusmn;\u0026thinsp;0.95 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.17\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e24.82\u0026thinsp;\u0026plusmn;\u0026thinsp;2.35 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e18.05\u0026thinsp;\u0026plusmn;\u0026thinsp;1.05 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ca\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCAT\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.91\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 Ab\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003ePOD\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ca\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.62\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.16\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003ePPO\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.28\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Ca\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003ePeroxide\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e70.3\u0026thinsp;\u0026plusmn;\u0026thinsp;17.1 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e65.5\u0026thinsp;\u0026plusmn;\u0026thinsp;6.0 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e29.7\u0026thinsp;\u0026plusmn;\u0026thinsp;3.2 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e53.7\u0026thinsp;\u0026plusmn;\u0026thinsp;12.7 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e43.0\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e27.4\u0026thinsp;\u0026plusmn;\u0026thinsp;2.8 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSuperoxide\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e22.50\u0026thinsp;\u0026plusmn;\u0026thinsp;6.30 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e21.41\u0026thinsp;\u0026plusmn;\u0026thinsp;3.47 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.73\u0026thinsp;\u0026plusmn;\u0026thinsp;1.47 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e32.20\u0026thinsp;\u0026plusmn;\u0026thinsp;2.92 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e15.52\u0026thinsp;\u0026plusmn;\u0026thinsp;4.26 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e6.67\u0026thinsp;\u0026plusmn;\u0026thinsp;1.38 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eMDA\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.34\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.89\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003eGas exchange and chlorophyll\u003c/b\u003e \u003cb\u003ea\u003c/b\u003e \u003cb\u003eresponses\u003c/b\u003e\u003c/p\u003e\u003cp\u003eHigher light intensities significantly reduced net CO\u003csub\u003e2\u003c/sub\u003e assimilation (\u003cem\u003eA\u003c/em\u003e), transpiration (\u003cem\u003eE\u003c/em\u003e), stomatal conductance (\u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e), and carboxylation efficiency (\u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). WUE increased with light intensity across all treatments, regardless of fertilization. In the control group, WUE\u003csub\u003eint\u003c/sub\u003e followed the same pattern, while fertilized plants showed the opposite trend, with a peak under VLR. Fertilized plants showed reduced \u003cem\u003eA\u003c/em\u003e, \u003cem\u003eE\u003c/em\u003e, \u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e, and \u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e compared to controls, except for \u003cem\u003eA\u003c/em\u003e under VLR and for \u003cem\u003eE\u003c/em\u003e and \u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e under HR, where differences were not significant. WUE and WUE\u003csub\u003eint\u003c/sub\u003e were higher in fertilized plants under VLR but declined under HR, reversing the initial advantage. \u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e remained higher in fertilized plants under VLR, while under HR, control plants surpassed them. Figure\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e presents the fluorescence parameters across treatments. Overall, F\u003csub\u003ev\u003c/sub\u003e\u0026rsquo;/F\u003csub\u003em\u003c/sub\u003e\u0026rsquo; was lower in plants under HR compared to other light conditions. Φ\u003csub\u003ePSII\u003c/sub\u003e and ETR followed a similar pattern, with the highest values under VLR, regardless of fertilization. Comparing control and manure-fertilized plants, F\u003csub\u003ev\u003c/sub\u003e\u0026rsquo;/F\u003csub\u003em\u003c/sub\u003e\u0026rsquo; was significantly higher in the control group, while Φ\u003csub\u003ePSII\u003c/sub\u003e, \u003cem\u003eq\u003c/em\u003e\u003csub\u003ep\u003c/sub\u003e, and ETR were consistently higher in fertilized plants across light intensities. The exception was under HR, where Φ\u003csub\u003ePSII\u003c/sub\u003e and ETR showed no significant differences between groups.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eGas exchanges under dehydration and after rehydration\u003c/h2\u003e\u003cp\u003eThe most pronounced changes in gas exchange occurred across dehydration durations within each light environment (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Under VLR, \u003cem\u003eA\u003c/em\u003e remained stable after 10 days but declined significantly after 20 days. Under HR, \u003cem\u003eA\u003c/em\u003e dropped more sharply, especially after 20 days. \u003cem\u003eE\u003c/em\u003e followed a similar trend, with greater reductions under HR. \u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e decreased significantly in both light conditions after 20 days, with a more pronounced effect under HR. WUE and WUE\u003csub\u003eint\u003c/sub\u003e means varied with dehydration time but showed no significant differences between treatments. \u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e remained unchanged, except after 20 days in plants under HR, when values were lower. Following rehydration, differences among dehydration treatments diminished, and values of \u003cem\u003eA\u003c/em\u003e, \u003cem\u003eE\u003c/em\u003e, \u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e, and \u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e increased, particularly under HR, where the recovery was statistically significant. During the experiment, a significant decline in soil moisture was observed after 10 and 20 days of dehydration under both VLR and HR conditions (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). Plant water potential also decreased with dehydration, particularly under HR, dropping from \u0026minus;\u0026thinsp;0.20 MPa in the control to -4.63 MPa after 20 days (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Following rehydration, plant water potential showed marked recovery, with no significant differences among treatments. Comparing plants subjected to 20 days of dehydration before and after rehydration, the increase in water potential was significant under HR.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eGas exchange parameters in \u003cem\u003eSyzygium jambos\u003c/em\u003e plants subjected to different dehydration durations (10 and 20 days) under very low radiation (VLR) and high radiation (HR), before and after rehydration. Parameters include CO\u003csub\u003e2\u003c/sub\u003e assimilation rate (\u003cem\u003eA\u003c/em\u003e, \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), transpiration rate (\u003cem\u003eE\u003c/em\u003e, mmol H\u003csub\u003e2\u003c/sub\u003eO m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), stomatal conductance (\u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e, mol H\u003csub\u003e2\u003c/sub\u003eO m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), intracellular-to-ambient CO\u003csub\u003e2\u003c/sub\u003e ratio (\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ea\u003c/sub\u003e), water-use efficiency (WUE, \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e mmol\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e H\u003csub\u003e2\u003c/sub\u003eO), intrinsic water use efficiency (WUE\u003csub\u003eint\u003c/sub\u003e, \u0026micro;mol CO\u003csub\u003e2\u003c/sub\u003e mol\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e H\u003csub\u003e2\u003c/sub\u003eO), and carboxylation efficiency (\u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e, \u0026micro;mol m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e Pa\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Uppercase letters indicate comparisons of dehydration duration within each light condition, while lowercase letters compare the effects of light conditions within each dehydration treatment. Data are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability. Asterisks denote statistically significant differences between dehydration and post-rehydration values (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"9\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"7\" nameend=\"c9\" namest=\"c3\"\u003e\u003cp\u003eUnder dehydration\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eParameter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003econtrol\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10 days\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e20 days\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003econtrol\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e10 days\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e20 days\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eA\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e5.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.30 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e2.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Ca\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eE\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.19\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.98\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.98\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00 Ca\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eg\u003c/b\u003e\u003csub\u003e\u003cb\u003es\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.097\u0026thinsp;\u0026plusmn;\u0026thinsp;0.013 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.072\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.004\u0026thinsp;\u0026plusmn;\u0026thinsp;0.000 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.050\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.018\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004 ABb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.001\u0026thinsp;\u0026plusmn;\u0026thinsp;0.000 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eC\u003c/b\u003e\u003csub\u003e\u003cb\u003ei\u003c/b\u003e\u003c/sub\u003e\u003cb\u003e/\u003c/b\u003e\u003cb\u003eC\u003c/b\u003e\u003csub\u003e\u003cb\u003ea\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 ABa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eWUE\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.59\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8.59\u0026thinsp;\u0026plusmn;\u0026thinsp;2.19 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e5.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e6.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.76 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e7.18\u0026thinsp;\u0026plusmn;\u0026thinsp;1.55 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eWUE\u003c/b\u003e\u003csub\u003e\u003cb\u003eint\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e54.0\u0026thinsp;\u0026plusmn;\u0026thinsp;7.3 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e63.6\u0026thinsp;\u0026plusmn;\u0026thinsp;7.5 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e162.1\u0026thinsp;\u0026plusmn;\u0026thinsp;41.0 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e101.8\u0026thinsp;\u0026plusmn;\u0026thinsp;1.8 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e137.6\u0026thinsp;\u0026plusmn;\u0026thinsp;17.0 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e153.9\u0026thinsp;\u0026plusmn;\u0026thinsp;31.4 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eA\u003c/b\u003e\u003cb\u003e/\u003c/b\u003e\u003cb\u003eC\u003c/b\u003e\u003csub\u003e\u003cb\u003ei\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.017\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.016\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.008\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.023\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.015\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.001\u0026thinsp;\u0026plusmn;\u0026thinsp;0.000 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"7\" nameend=\"c9\" namest=\"c3\"\u003e\u003cp\u003e\u003cb\u003eAfter rehydration\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eA\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.90\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.68\u0026thinsp;\u0026plusmn;\u0026thinsp;0.62 Aa*\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.40 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e3.18\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e3.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.44 Aa*\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eE\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.18\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.73\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e1.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.76\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Aa*\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eg\u003c/b\u003e\u003csub\u003e\u003cb\u003es\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.097\u0026thinsp;\u0026plusmn;\u0026thinsp;0.013 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.063\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004 ABa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.042\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.075\u0026thinsp;\u0026plusmn;\u0026thinsp;0.018 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.032\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.040\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003 Aa*\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eC\u003c/b\u003e\u003csub\u003e\u003cb\u003ei\u003c/b\u003e\u003c/sub\u003e\u003cb\u003e/\u003c/b\u003e\u003cb\u003eC\u003c/b\u003e\u003csub\u003e\u003cb\u003ea\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.77\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.59\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eWUE\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.24\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.50\u0026thinsp;\u0026plusmn;\u0026thinsp;0.16 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.53 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4.71\u0026thinsp;\u0026plusmn;\u0026thinsp;0.53 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e5.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.73 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e4.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eWUE\u003c/b\u003e\u003csub\u003e\u003cb\u003eint\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e52.9\u0026thinsp;\u0026plusmn;\u0026thinsp;8.2 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e77.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e94.9\u0026thinsp;\u0026plusmn;\u0026thinsp;14.1 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e69.5\u0026thinsp;\u0026plusmn;\u0026thinsp;10.9 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e105.1\u0026thinsp;\u0026plusmn;\u0026thinsp;13.9 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e91.5\u0026thinsp;\u0026plusmn;\u0026thinsp;5.4 Aa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eA\u003c/b\u003e\u003cb\u003e/\u003c/b\u003e\u003cb\u003eC\u003c/b\u003e\u003csub\u003e\u003cb\u003ei\u003c/b\u003e\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.016\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.018\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.016\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.017\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.015\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.015\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002 Aa*\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSoil temperature, soil moisture, and leaf water potential (Ψw) of \u003cem\u003eSyzygium jambos\u003c/em\u003e plants subjected to dehydration for 10 and 20 days under very low radiation (VLR) and high radiation (HR), followed by rehydration. Uppercase letters indicate comparisons of dehydration duration within each light condition, while lowercase letters compare the effects of light conditions within each dehydration treatment. Data are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard error. Means followed by the same letters do not differ significantly according to the Scott-Knott test at a 5% probability. Asterisks denote statistically significant differences between dehydration and post-rehydration values (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"9\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"7\" nameend=\"c9\" namest=\"c3\"\u003e\u003cp\u003eUnder dehydration\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e\u003cp\u003eVLR\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"3\" nameend=\"c9\" namest=\"c7\"\u003e\u003cp\u003eHR\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eParameter\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003econtrol\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10 days\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e20 days\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003econtrol\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003e10 days\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003e20 days\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSoil temperature (\u0026deg;C)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e21.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e25.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e25.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e27.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e33.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e29.8\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5 ABa\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSoil moisture (%)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e8.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e3.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5 Ab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e0.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 Ba\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eΨw (MPa)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-0.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-1.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.54 Ba\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-0.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-1.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25 Bb\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-4.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 Cb\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"7\" nameend=\"c9\" namest=\"c3\"\u003e\u003cp\u003eAfter rehydration\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eΨw (MPa)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-0.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-0.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-0.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-0.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 Aa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003e-0.20\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01 Aa*\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eSoil pH, or active acidity, reflects H\u003csup\u003e+\u003c/sup\u003e concentration in solutions. Based on classification of Soil Survey Staff (\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), the control treatment had pH within the \u0026ldquo;neutral\u0026rdquo; range (6.6\u0026ndash;7.3), the bovine manure treatment had pH values within the \u0026ldquo;slightly acid\u0026rdquo; range (6.1\u0026ndash;6.5), with the manure-amended substrate within the ideal range for plant growth (5.5\u0026ndash;6.5), at pH 6.2 (Islam et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1980\u003c/span\u003e). The chemically fertilized substrate falls within the \u0026ldquo;very strong acid\u0026rdquo; range (4.5-5.0), nearing the \u0026ldquo;extremely high\u0026rdquo; threshold (3.5\u0026ndash;4.4). Excess acidity can increase the solubility of toxic elements, such as manganese and aluminum, which impair root function and cell development (Kochian \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Horst et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). This pH condition can potentially reduce yield, which may have been the primary factor in the death of plants observed in treatments supplemented with chemical fertilizer. Organic amendment (with bovine manure) enriched the substrates with nutrients and improved chemical attributes by increasing pH and base saturation while reducing acidity. The absence of Al indicates that the amendment effectively buffered soil acidity, preventing Al mobilization. Over the course of the experiment, pH values became more basic in fertilized substrates. This shift is likely linked to the decomposition of manure, which releases carbonates, bicarbonates, and organic acids \u0026ndash; compounds that buffer acidity and raise substrate pH (Whalen et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Holatko et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eBefore planting, manure-amended substrates contained higher levels of most macro- and micronutrients than the control (except Fe and Cu). This pattern largely persisted after cultivation for K, P, Ca, Mg, OM, SB, t, T, and V across all light environments. Thus, while manure improved substrate fertility and buffering capacity, the intensity of radiation determined the rate and direction of nutrient transformations in the soil. Over the 11-month experimental period, nutrient depletion was more pronounced in manure-amended substrates under HR, especially for K, P, Mg, Cu, B and S, as well as for the parameters SB and T, reflecting greater nutrient demand for plant growth.\u003c/p\u003e\u003cp\u003eFoliar N content decreased with increasing light intensity in both control and fertilized plants, consistent with findings in \u003cem\u003eSinarundinaria nitida\u003c/em\u003e, where shaded plants maintained higher N and chlorophyll levels (Yang et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2014b\u003c/span\u003e). In the control group, P, K, Ca, Mg, B, and Mn remained relatively stable across light environments, while in the manure-treated group, nutrient levels fluctuated without a clear pattern. Fertilized plants accumulated, under high radiation, more P, Mg, Cu, B, and Zn, whereas K, Ca, and S decreased. These variable responses align with reports of light-dependent nutrient allocation in other tropical tree species, such as \u003cem\u003eSwietenia macrophylla\u003c/em\u003e and \u003cem\u003eDipteryx odorata\u003c/em\u003e (Gon\u0026ccedil;alves et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), and with studies noting inconsistent effects of light intensity on macronutrient composition (Ronquim et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Dalmolin et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThese patterns in nutrient dynamics also help explain the observed physiological responses. Plants under HR exhibited higher total protein content than those under VLR and LR, regardless of fertilization. Interestingly, despite lower foliar N levels under HR, protein content was higher. This apparent discrepancy reflects the complex distribution of nitrogen within leaf tissues, which can shift between photosynthetic and structural pools depending on environmental conditions (Wang et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). According to these authors, foliar nitrogen is allocated among several functional compartments, including photosynthetic proteins such as RuBisCo and components of the light-harvesting complexes, as well as non-photosynthetic proteins in cell walls, mitochondria, and cytosol. \u003cem\u003eS. jambos\u003c/em\u003e also exhibited higher chlorophyll concentrations under VLR, as confirmed by spectrophotometric analysis and by non-destructive analyses with SPAD-502 (Resende et al. unpublished data), suggesting a greater investment in light-harvesting structures under low-light conditions. This allocation pattern likely explains the higher foliar N levels observed under VLR despite lower total protein content, as more nitrogen was directed to chlorophyll and associated proteins rather than to bulk protein synthesis. Positive correlations between foliar N and chlorophyll content have also been reported in other species (Mizusaki et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2014a\u003c/span\u003e; Schlichting et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eProline levels exhibited divergent responses: they increased under HR in the control group but decreased under LR and HR in organic fertilizer plants. Proline accumulation differs with fertilization and light, suggesting that its role in osmoprotection or stress mitigation varies depending on nutrient status. Although proline stabilizes subcellular structures and acts as a redox buffer (Reddy et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Molinari et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Verslues and Sharma \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), its primary role is to facilitate osmotic adjustment under water stress (Hong et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Carvalho et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eSOD serves as the first enzymatic barrier against ROS, converting O\u003csub\u003e2\u003c/sub\u003e.\u003csup\u003e-\u003c/sup\u003e to H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e (Alscher et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), which is then scavenged by CAT and various PODs to prevent the formation of highly reactive species, such as OH. (Kibinza et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Shivashankara et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). PPO, abundant in plants, oxidizes phenolics into quinones and may act synergistically with POD (Krishna et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Overall, SOD and PPO activities were significantly lower under HR, as were H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e and superoxide levels. Such reductions in SOD and PPO under HR may reflect enzyme inhibition or a strategic shift in antioxidant defense, with other enzymes (POD, CAT) compensating depending on light intensity. CAT and POD showed complementary activity: CAT was more active under VLR and HR, while POD peaked under LR. Lipid peroxidation (TBARs index) was highest under HR.\u003c/p\u003e\u003cp\u003eThese patterns indicate that high radiation may trigger protein synthesis and lipid peroxidation as part of a protective response to light stress, while simultaneously reducing ROS accumulation through efficient scavenging mechanisms. This finding aligns with the observed increase in lipid peroxidation but lower ROS levels under HR, suggesting a dynamic regulation of defense pathways. These results also indicate increased ROS production in shaded environments (LR) and an effective response to oxidative stress under higher radiation. Interestingly, lower H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e and superoxide levels under HR may reflect alternative antioxidant defenses, such as carotenoids, which deactivate singlet oxygen and other ROS. Indeed, \u003cem\u003eS. jambos\u003c/em\u003e under HR exhibited significantly lower chlorophyll/carotenoid ratios (Resende et al. unpublished), highlighting the photoprotective role of these pigments in full sunlight conditions. Moreover, fertilization modulates these stress responses, likely by influencing nutrient-dependent metabolic pathways and antioxidant capacity.\u003c/p\u003e\u003cp\u003eContrasting results are reported in the literature regarding these enzymes activities. Yang et al. (\u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) found increased SOD and CAT activities, and reduced POD activity in \u003cem\u003ePicea asperata\u003c/em\u003e under high light conditions. The role of CAT in H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e detoxification under intense radiation was demonstrated in \u003cem\u003eArabidopsis thaliana\u003c/em\u003e mutants, which exhibited elevated H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e levels and cell death (Vandenabeele et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Vanderauwera et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Liu et al. (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) observed significantly higher levels of hydrogen peroxide and superoxide radicals, along with increased activities of CAT, SOD, and APX, in \u003cem\u003eAlnus\u003c/em\u003e species grown under full sunlight, with no significant changes in lipid peroxidation. In \u003cem\u003eTorreya grandis\u003c/em\u003e, SOD activity was light-dependent, peaking at both the extremes of light levels, indicating that both excessive and insufficient light can impair plant metabolism (Tang et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn the present study, gas exchange measurements were conducted under a standardized light intensity (1,000 \u0026micro;mol photons m\u003csup\u003e-2\u003c/sup\u003e s\u003csup\u003e-1\u003c/sup\u003e) using the IRGA LED light chamber. The higher values of \u003cem\u003eA\u003c/em\u003e, E, \u003cem\u003egs\u003c/em\u003e, and \u003cem\u003eA\u003c/em\u003e/Ci observed in plants from shaded environments indicate an enhanced capacity for light utilization under limited irradiance. Furthermore, these results demonstrate the absence of photoinhibition, enabling shade-adapted plants to efficiently exploit the available light in environments where this resource is scarce. Shade plants in forest understories often experience brief, high-intensity sunflecks throughout the day, which contribute substantially to their daily carbon gain. These species are able to utilize sunflecks for positive net photosynthesis without suffering significant photoinhibition, due to rapid activation of energy dissipation mechanisms and other physiological adaptations (Way and Pearcy \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, Zhang et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eIn contrast, elevated WUE in plants exposed to HR reflects efficient carbon assimilation relative to transpiration, indicating reduced water loss. Different patterns were reported for \u003cem\u003ePiper aduncum\u003c/em\u003e, a pioneer species, which showed increased \u003cem\u003eA\u003c/em\u003e and \u003cem\u003eE\u003c/em\u003e and reduced WUE under higher light (Pacheco et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Similarly, \u003cem\u003eAlnus formosana\u003c/em\u003e, an invasive species, exhibited higher \u003cem\u003eA\u003c/em\u003e, \u003cem\u003eE\u003c/em\u003e, and \u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e under high radiation (Liu et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Compared to the native species \u003cem\u003eAlnus cremastogyne\u003c/em\u003e, \u003cem\u003eA. formosana\u003c/em\u003e demonstrated superior photosynthetic and stomatal performance, highlighting functional advantages linked to invasiveness. In our study, high irradiance led to reduced net photosynthesis and increased stomatal limitation, reducing CO\u003csub\u003e2\u003c/sub\u003e assimilation and carboxylation efficiency.\u003c/p\u003e\u003cp\u003eIn contrast to our work, Silva et al. (\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) reported increased RuBisCo carboxylation efficiency (\u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e) under high light, suggesting species-specific responses to irradiance. These findings underscore the complexity of light-mediated physiological responses and the importance of ecological strategy in shaping gas exchange dynamics. Photosynthetic traits of plants from different successional stages were compared under contrasting light conditions (Zhang et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Pioneer species showed greater reductions in photosynthetic rates under shade compared to mid- and late-successional species. This reflects the generally lower shade tolerance of early successional species and their reliance on open (under high light) environments for optimal carbon gain. Additional traits \u0026ndash; such as lower light compensation points, reduced respiration rates, and higher apparent quantum yield \u0026ndash; suggest that late-successional species are better adapted to low-light conditions. This response aligns with our findings, considering that light levels in the understory of mature forests, where these species grow and \u003cem\u003eS. jambos\u003c/em\u003e is commonly found, can drop below 2% of full daylight (Chazdon and Fetcher \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). Such environments favor species with enhanced light-use efficiency and physiological plasticity.\u003c/p\u003e\u003cp\u003eThe ΦPSII represents the fraction of absorbed light energy allocated to photochemical processes and typically shows higher values in plants exposed to low irradiance (Yang et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Species adapted to high light intensities tend to exhibit lower effective quantum yield of PSII, as a larger portion of absorbed light is dissipated through non-photochemical pathways. This strategy helps to protect the photosynthetic apparatus from excess energy, minimizing photoinhibition (Yang et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). This pattern was confirmed in our study, where measured fluorescence parameters were generally higher in plants grown under VLR compared to those under HR. Previous studies support these findings. Zhang et al. (\u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), for example, reported increased ΦPSII in response to reduced light intensity in five tree species; for \u003cem\u003eQuercus aliena\u003c/em\u003e, shaded plants exhibited higher ΦPSII and ETR values (Xu et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Enhanced photochemical efficiency under low light likely contributes to maintaining positive carbon balance in shaded environments, reinforcing the ecological advantage of high shade tolerance in \u003cem\u003eS. jambos\u003c/em\u003e.\u003c/p\u003e\u003cp\u003eVariations in gas exchange parameters across dehydration durations and light environments reveal a nuanced interaction between water availability and irradiance. Under HR, sharper declines in \u003cem\u003eA\u003c/em\u003e, \u003cem\u003eE\u003c/em\u003e, and \u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e after 20 days of dehydration reflect intensified physiological stress. Stomatal closure under drought is a primary adaptive response to minimize transpirational water loss. However, this response also restricts CO\u003csub\u003e2\u003c/sub\u003e uptake, thereby limiting photosynthetic carbon assimilation (Flexas et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2004\u003c/span\u003e, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, Oguz et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Despite these reductions, the stability of WUE suggests a proportional decline in both CO\u003csub\u003e2\u003c/sub\u003e assimilation and water loss, reflecting balanced stomatal regulation. While stomatal closure explains the initial reductions in gas exchange, the observed drop in \u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e after 20 days of dehydration under HR points to additional non-stomatal limitations, likely linked to biochemical constraints on photosynthesis (Flexas and Medrano \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Moreover, the overall stability of \u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e suggests that such biochemical limitations only emerged under more severe stress, highlighting the species\u0026rsquo; resilience to combined environmental pressures. Under high irradiance, the compounded effects of water deficit and excess energy intensify photoprotective demands, leading to increased non-photochemical energy dissipation and further constraining carbon assimilation (Nosalewicz et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eOur findings align with recent work by He et al. (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), who reported similar patterns in tropical perennial trees: \u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e, \u003cem\u003eA\u003c/em\u003e, and \u003cem\u003eE\u003c/em\u003e d ropped significantly under drought, followed by strong recovery after rehydration. In our study, the substantial recovery of \u003cem\u003eA\u003c/em\u003e, \u003cem\u003eE\u003c/em\u003e, \u003cem\u003eg\u003c/em\u003e\u003csub\u003es\u003c/sub\u003e, and \u003cem\u003eA\u003c/em\u003e/\u003cem\u003eC\u003c/em\u003e\u003csub\u003ei\u003c/sub\u003e \u0026ndash; especially under HR \u0026ndash; along with the rapid restoration of gas exchange and water potential after rehydration, reveals the strong desiccation tolerance and hydraulic recovery mechanisms of \u003cem\u003eS. jambos\u003c/em\u003e. The rebound in plant water potential, particularly under HR, further supports the species\u0026rsquo; resilience to episodic drought, provided rehydration occurs. Such resilience and plasticity under fluctuating environmental conditions may help explain the invasive potential of \u003cem\u003eS. jambos\u003c/em\u003e, especially in regions prone to intermittent drought and high irradiance.\u003c/p\u003e\u003cp\u003eThe results of our study allow us to conclude that \u003cem\u003eS. jambos\u003c/em\u003e exhibits remarkable physiological and biochemical plasticity, enabling it to thrive under varying light intensities, fertilization regimes, and water availability. Light intensity strongly influenced nitrogen allocation, protein synthesis, antioxidant activity, and gas exchange. The species maintains photosynthetic efficiency and photochemical performance across contrasting light environments, deploys effective antioxidant and osmotic stress responses, and recovers rapidly from dehydration. Its flexible allocation of nitrogen and pigments, combined with tolerance to both shade and high irradiance, underpins its ecological versatility and likely contributes to its invasive potential. Understanding these adaptive mechanisms is essential for predicting their behavior under climate change and informing environmental management strategies.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors thank the Funda\u0026ccedil;\u0026atilde;o de Amparo \u0026agrave; Pesquisa do Estado de Minas Gerais (FAPEMIG) and the Coordena\u0026ccedil;\u0026atilde;o de Aperfei\u0026ccedil;oamento de Pessoal de N\u0026iacute;vel Superior (CAPES) for their financial support.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBoth authors contributed to the conception and design of the study. Material preparation, data collection, and analysis were performed by Cristiano Ferrara de Resende. The first draft of the manuscript was written by Cristiano Ferrara de Resende and both authors commented on previous versions. Both authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by FAPEMIG (Project CRA-APQ 00974/13). Author C.F.R. has received a doctoral scholarship from CAPES.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eData will be made available from the corresponding author on request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAdelino, J. R. P., Heringer, G., Diagne, C., Courchamp, F., Faria, L. D. B., \u0026amp; Zenni, R. D. (2021). 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Effects of water stress on chlorophyll fluorescence and growth of \u003cem\u003eRorippa amphibia\u003c/em\u003e: a well-adjusted invasive plant in China. \u003cem\u003eActa Physiologiae Plantarum\u003c/em\u003e, 46, 97. https://doi.org/10.1007/s11738-024-03722-z\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"invasive species, photosynthesis, chlorophyll a fluorescence, water stress, light acclimation","lastPublishedDoi":"10.21203/rs.3.rs-8196016/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8196016/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003ePurpose\u003c/h2\u003e\u003cp\u003eThis study explores the physiological and biochemical plasticity of \u003cem\u003eSyzygium jambos\u003c/em\u003e, an invasive tree species native to Tropical Asia, under contrasting light intensities, fertilization regimes, and water availability. The aim is to elucidate the mechanisms underlying its ecological success and to address how these mechanisms may inform strategies for managing invasive plants under changing climate scenarios.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e\u003cp\u003eA factorial experiment was conducted using seedlings grown in two substrates \u0026ndash; control (soil and sand), and organic amendment (aged bovine manure) \u0026ndash; and exposed them to three light environments: very low radiation (VLR), low radiation (LR), and high radiation (HR). Experiments were conducted under controlled conditions at the Federal University of Juiz de Fora (Minas Gerais, Brazil). Physiological assessments included gas exchange, chlorophyll \u003cem\u003ea\u003c/em\u003e fluorescence, antioxidant enzyme activities, osmotic stress markers, and foliar nutrient profiling.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e\u003cp\u003ePlants under HR exhibited elevated protein content and lipid peroxidation levels, alongside reduced superoxide dismutase (SOD) and polyphenol oxidase (PPO) activities. Organic fertilization enhanced nutrient availability and modulated stress responses. Despite reduced photosynthetic rates under HR, \u003cem\u003eS. jambos\u003c/em\u003e maintained photochemical efficiency and water-use balance, with rapid recovery following drought stress. Nutrient analyses revealed significant differences between control and fertilized plants, with fertilized seedlings showing elevated levels of nitrogen (N), phosphorus (P), and potassium (K).\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e\u003cp\u003eThese findings reveal that \u003cem\u003eS. jambos\u003c/em\u003e possesses high ecophysiological plasticity, enabling it to adapt to heterogeneous tropical environments and recover from abiotic stress. Such versatility likely contributes to its invasive potential and ecological dominance. By linking functional traits to invasion success, this study provides a predictive framework for managing exotic species and assessing their impact under climate variability.\u003c/p\u003e","manuscriptTitle":"Physiological and biochemical plasticity as a driver of invasiveness in Syzygium jambos (L.) 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