
Figure 1.
F. hybrida 'White River' (left) and experimental layout in soil-based greenhouse (right).
Table 1.
Microbial traits and compositions of PGPB formulations implemented in the trial.
| Code | Characteristics | Active components | Proportions (CFU · mL-1) |
|---|---|---|---|
| A1 | Siderophores production | Bacillus toyonensis, Lysinibacillus fusiformis | 1 × 107 |
| A2 | IAA production | Bacillus toyonensis, Pseudomonas putida | 1 × 107 |
| A3 | Phosphate solubilisation | Bacillus proteolyticus, Pseudomonas batumici | 1 × 107 |
| A4 | Potassium solubilisation | Pseudomonas lini, Bacillus safensis | 1 × 107 |
| A5 | N2 fixer | Pseudomonas konensis, Bacillus thuringiensis | 1 × 107 |

Figure 2.
Daily mean temperature (red) and relative humidity (blue, dashed) in the greenhouse, shown with dual y-axes.
Table 2.
Soil properties by treatment.
| Grp. | PH | CaCO. (%) | Salt (%) | Sat (%) | OM (%) | Ntotal (%) | P avail | Kavail | Caextr | Mgextr | Feextr | Mnextr | Znextr | Cuextr |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Bad | 5.53 ± 0.05 c | 0.86 ± 0.11b | 0.15 ±0.00 a | 75.6 ± 0.57 b | 3.18 ± 0.02 c | 0.21 ± 0.00 b | 26.4 ± 0.59 c | 89.4 ± 1.22 a | 2284 ± 19.68 ab | 282 ± 1.80 b | 33.3 ± 1.10 b | 15.9 ± 0.40 b | 1.91 ± 0.05 c | 1.92 ± 0.07 c |
| C.F. | 6.22 ±0.00 a | 1.70 ± 0.2 a | 0.04 ±0.001 c | 78.6 ±0.57 a | 3.65 ± 0.01b | 0.23 ± 0.00 a | 43.52 ±2.87 a | 83.3 ± 1.85 b | 2184 ± 77.86 b | 233 ± 7.90 c | 33.7 ± 0.80 b | 8.10 ±0.21 c | 2.64 ± 0.08 b | 2.21 ± 0.07 b |
| Cont | 5.83 ± 0.05 b | 0.83 ± 0.05 b | 0.10 ± 0.0005 b | 77.6 ± 1.154 a | 4.25 ± 0.025 a | 0.20 ± 0.00 c | 39.7 ± 1.086 b | 73.5 ± 1.80 c | 2341 ± 37.06 a | 316 ± 7.29 a | 37.4 ± 0.75 a | 15.4 ± 0.22 b | 2.79 ± 0.05 a | 3.87 ±0.12 a |
| LSD | 0.09 | 0.27 | 0.00 | 1.63 | 0.04 | 0.00 | 3.61 | 3.30 | 102.05 | 12.59 | 1.80 | 0.59 | 0.13 | 0.19 |
| %CV | 0.80 | 12.12 | 0.11 | 1.06 | 0.54 | 1.22 | 4.93 | 2.01 | 2.25 | 2.27 | 2.59 | 2.16 | 2.68 | 3.56 |
Table 3.
Effects of PGPB consortia and chemical fertilisation on bulb development and reproductive traits in F. hybrida.
| Grp. | CDW (g · parcel-1) | Net Δ CFW (g · parcel-1) | Net Δ CD (mm · parcel-1) | NC (number · parcel-1) | LA (cm2) | CHL |
|---|---|---|---|---|---|---|
| A1 | 3.47 ± 0.26 c | 8.87 ± 1.33 b | 16.13 ± 0.67 c | 3.13 ± 0.34 a | 85.42 ± 10.29 bc | 59.58 ± 6.52 a |
| A2 | 4.20 ± 0.42 a | 10.46 ± 0.36 a | 21.15 ± 1.58 a | 2.53 ± 0.50 bc | 91.11 ± 9.65 a | 60.35 ± 5.65 a |
| A3 | 3.15 ± 0.51 d | 9.21 ± 1.33 b | 15.04 ± 1.24 d | 2.17 ± 0.38 d | 86.59 ± 11.81 bc | 58.03 ± 7.12 ab |
| A4 | 3.94 ± 0.49 b | 10.51 ± 0.31 a | 17.35 ± 1.59 b | 2.97 ± 0.14 a | 87.04 ± 9.90 ab | 59.64 ± 6.08 a |
| A5 | 3.27 ± 0.50 d | 8.37 ± 1.25 c | 15.09 ± 1.88 d | 2.62 ± 0.49 b | 85.09 ± 9.95 bc | 55.26 ± 8.11 b |
| C.F. | 3.21 ± 0.48 d | 9.03 ± 1.27 b | 14.10 ± 1.67 e | 2.40 ± 0.49 c | 84.90 ± 10.04 bc | 56.32 ± 4.22 b |
| C | 3.28 ± 0.54 d | 8.30 ± 0.97 c | 14.37 ± 2.01 e | 2.00 ± 0.21 e | 82.55 ± 9.56 c | 52.19 ± 8.37 c |
| LSD | 0.29 | 0.77 | 0.66 | 0.16 | 4.23 | 4.92 |
| %CV | 12.33 | 11.52 | 9.77 | 15.38 | 11.83 | 11.72 |
1 C, control; C.F., chemical fertiliser; CDW, corm dry weight (g · parcel-1); CFW, corm fresh weight; CHL, chlorophyll content (SPAD units; chlorophyll index); CV, coefficient of variation (%); LA, leaf area (cm2); LSD, least significant difference at p < 0.05; NC, number of cormlets (number · parcel-1); Net Δ CD, net change in corm diameter (mm · parcel-1); Net Δ CFW, net change in corm fresh weight (g · parcel-1); PGPB, plant growth-promoting bacteria; PGPR, plant growth-promoting rhizobacteria.

Figure 3.
Representative corms of F. hybrida under PGPR treatments (A1–A5), fertiliser (C.F.) and control (C). Within each panel: PGPR (left), fertiliser (middle), control (right). C.F., chemical fertilizer; PGPR, plant growth-promoting rhizobacteria.
Table 4.
Bulb content analyses after harvest.
| Grp. | N | P | K | Ca | Mg | Fe | Mn | Zn | Cu |
|---|---|---|---|---|---|---|---|---|---|
| A1 | 1.85 ±0.011 e | 0.19 ± 0.03 b | 0.63 ± 0.032 e | 0.28 ± 0.005 b | 0.12 ± 0.005 d | 1362 ± 33.45 e | 27.4 ± 1.153 cd | 10.03 ± 1.87 c | 4.23 ± 0.28 e |
| A2 | 2.10 ± 0.015 b | 0.24 ±0.00 a | 0.76 ± 0.015 b | 0.32 ±0.005 a | 0.25 ±0.005 a | 4660 ±61.46 a | 83.53 ±2.122 a | 18.36 ±0.28 a | 11.93 ±0.11 a |
| A3 | 1.93 ± 0.017 d | 0.14 ± 0.00 d | 0.64 ± 0.05 de | 0.27 ± 0.005 b | 0.10 ±0.01 e | 394 ± 20.54 f | 26.1 ± 19.52 de | 4.80 ±0.2 e | 3.73 ±0.51 f |
| A4 | 2.35 ±0.005 a | 0.19 ± 0.00 b | 0.82 ±0.005 a | 0.23 ± 0.005 c | 0.15 ± 0.005 c | 2106 ± 24.2 c | 39.36 ± 1.001 c | 10.46 ± 0.50 c | 5.73 ± 0.15 c |
| A5 | 1.80 ±0.011 f | 0.20 ± 0.05 b | 0.66 ±0.01 d | 0.22 ±0.001 d | 0.10 ± 0.00 e | 358 ± 17.95 f | 13.2 ± 0.66 e | 7.53 ± 1.10 d | 4.86 ± 0.25 d |
| C.F. | 1.71 ± 0.015 g | 0.16 ±0.05 cd | 0.59 ± 0.00 f | 0.21 ±0.01 d | 0.12 ± 0.005 d | 1546 ± 61.20 d | 30.6 ± 1.17 cd | 5.66 ± 0.68 e | 4.16 ± 0.15 ef |
| C. | 1.98 ±0.01 c | 0.18 ±0.01 be | 0.72 ± 0.005 c | 0.21 ±0.00 d | 0.18 ± 0.005 b | 3307 ± 94.90 b | 57.53 ± 2.35 b | 12.9 ± 0.55 b | 7.46 ± 0.25 b |
| LSD | 0.02 | 0.01 | 0.01 | 0.01 | 0.01 | 64.5 | 13.16 | 1.135 | 0.48 |
| %CV | 0.66 | 7.87 | 2.18 | 2.46 | 4.14 | 2.65 | 18.94 | 9.18 | 4.59 |

Figure 4.
PCA of morphophysiological and nutritional traits of F. hybrida. PCA, principal component analysis.
Table 5.
PCA loadings of morphophysiological and nutritional traits in F. hybrida.
| Contribution of the observations (%) | |||||
|---|---|---|---|---|---|
| F1 | F2 | F3 | F4 | F5 | |
| A1 | 0.325 | 14.725 | 16.077 | 0.000 | 12.147 |
| A2 | 63.360 | 2.646 | 2.755 | 7.090 | 3.387 |
| A3 | 7.720 | 5.919 | 3.687 | 54.765 | 13.054 |
| A4 | 5.723 | 23.365 | 20.972 | 27.441 | 7.961 |
| A5 | 9.469 | 0.512 | 39.624 | 8.856 | 7.101 |
| C | 0.076 | 49.967 | 3.043 | 1.411 | 9.453 |
| C.F. | 13.327 | 2.865 | 13.841 | 0.437 | 46.897 |

Figure 5.
PCA biplot of fertiliser treatments based on morphophysiological and nutritional traits in F. hybrida. PCA, principal component analysis,
Table 6.
Pearson correlation matrix among morphophysiological and nutritional traits of F. hybrida.
| Variables | CDW | NC | Net Δ CD | LA | CHL | Net Δ CFW | N | P | K | Ca | Mg | Fe | Mn | Zn | Cu |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| CDW | 1 | 0.943 | 0.439 | 0.823 | 0.699 | 0.861 | 0.576 | 0.795 | 0.780 | 0.605 | 0.810 | 0.715 | 0.724 | 0.796 | 0.758 |
| NC | 0.943 | 1 | 0.339 | 0.930 | 0.730 | 0.801 | 0.455 | 0.807 | 0.623 | 0.810 | 0.796 | 0.674 | 0.710 | 0.793 | 0.803 |
| Net Δ CD | 0.439 | 0.339 | 1 | 0.130 | 0.438 | 0.360 | 0.209 | 0.372 | 0.171 | 0.234 | -0.045 | -0.130 | -0.157 | 0.079 | -0.142 |
| LA | 0.823 | 0.930 | 0.130 | 1 | 0.810 | 0.718 | 0.371 | 0.670 | 0.651 | 0.829 | 0.730 | 0.593 | 0.644 | 0.734 | 0.774 |
| Chl | 0.699 | 0.730 | 0.438 | 0.810 | 1 | 0.601 | 0.439 | 0.468 | 0.619 | 0.736 | 0.500 | 0.373 | 0.390 | 0.575 | 0.432 |
| Net Δ CFW | 0.861 | 0.801 | 0.360 | 0.718 | 0.601 | 1 | 0.817 | 0.407 | 0.663 | 0.541 | 0.567 | 0.471 | 0.492 | 0.444 | 0.484 |
| N | 0.576 | 0.455 | 0.209 | 0.371 | 0.439 | 0.817 | 1 | -0.001 | 0.458 | 0.325 | 0.439 | 0.433 | 0.425 | 0.254 | 0.234 |
| P | 0.795 | 0.807 | 0.372 | 0.670 | 0.468 | 0.407 | -0.001 | 1 | 0.543 | 0.504 | 0.744 | 0.661 | 0.670 | 0.854 | 0.802 |
| K | 0.780 | 0.623 | 0.171 | 0.651 | 0.619 | 0.663 | 0.458 | 0.543 | 1 | 0.177 | 0.664 | 0.602 | 0.591 | 0.658 | 0.610 |
| Ca | 0.605 | 0.810 | 0.234 | 0.829 | 0.736 | 0.541 | 0.325 | 0.504 | 0.177 | 1 | 0.530 | 0.405 | 0.462 | 0.545 | 0.550 |
| Mg | 0.810 | 0.796 | -0.045 | 0.730 | 0.500 | 0.567 | 0.439 | 0.744 | 0.664 | 0.530 | 1 | 0.982 | 0.988 | 0.959 | 0.960 |
| Fe | 0.715 | 0.674 | -0.130 | 0.593 | 0.373 | 0.471 | 0.433 | 0.661 | 0.602 | 0.405 | 0.982 | 1 | 0.996 | 0.924 | 0.920 |
| Mn | 0.724 | 0.710 | -0.157 | 0.644 | 0.390 | 0.492 | 0.425 | 0.670 | 0.591 | 0.462 | 0.988 | 0.996 | 1 | 0.926 | 0.944 |
| Zn | 0.796 | 0.793 | 0.079 | 0.734 | 0.575 | 0.444 | 0.254 | 0.854 | 0.658 | 0.545 | 0.959 | 0.924 | 0.926 | 1 | 0.941 |
| Cu | 0.758 | 0.803 | -0.142 | 0.774 | 0.432 | 0.484 | 0.234 | 0.802 | 0.610 | 0.550 | 0.960 | 0.920 | 0.944 | 0.941 | 1 |
1 CDW, corm dry weight; CFW, corm fresh weight; CHL, chlorophyll content (SPAD units; chlorophyll index); LA, leaf area; NC, number of cormlets. Bold values in Table 6 represent the diagonal self-correlation coefficients (r = 1.00) for each trait.
Table 7.
Variable contributions (%) to principal components (F1-F5) in F. hybrida.
| F1 | F2 | F3 | F4 | F5 | |
|---|---|---|---|---|---|
| CDW | 9.298 | 2.352 | 0.064 | 3.831 | 0.449 |
| NC | 9.300 | 1.773 | 1.692 | 0.867 | 0.763 |
| Net Δ CD | 0.443 | 26.367 | 7.287 | 21.948 | 12.173 |
| LA | 8.212 | 1.165 | 1.591 | 8.995 | 9.854 |
| Chl | 5.358 | 10.412 | 1.478 | 2.892 | 16.346 |
| Net Δ CFW | 5.896 | 10.063 | 10.705 | 0.010 | 0.544 |
| N | 2.722 | 6.975 | 40.434 | 0.672 | 7.444 |
| P | 6.629 | 0.617 | 17.751 | 9.799 | 1.459 |
| K | 5.794 | 0.236 | 4.714 | 14.823 | 40.051 |
| Ca | 4.974 | 3.663 | 7.127 | 34.386 | 3.819 |
| Mg | 8.991 | 5.203 | 0.636 | 0.033 | 1.037 |
| Fe | 7.508 | 9.385 | 2.431 | 0.360 | 3.295 |
| Mn | 7.833 | 9.133 | 1.842 | 0.014 | 2.750 |
| Zn | 8.700 | 4.472 | 1.790 | 1.142 | 0.011 |
| Cu | 8.342 | 8.183 | 0.457 | 0.226 | 0.004 |

Figure 6.
Proposed mechanisms by which Bacillus and Pseudomonas spp. enhance F. hybrida growth through VOCs, IAA, siderophores, phosphate solubilisation and ACC deaminase. ACC, 1-aminocyclopropane-1-carboxylate; IAA, indole-3-acetic acid; VOCs, volatile organic compounds.