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Seed germination, seedling growth performance, genetic stability and biochemical responses of papaya (Carica papaya L.) upon pre-sowing seed treatments Cover

Seed germination, seedling growth performance, genetic stability and biochemical responses of papaya (Carica papaya L.) upon pre-sowing seed treatments

Open Access
|Mar 2025

Figures & Tables

Figure 1.

Effect of papaya seed pre-sowing treatments on seed GP. Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. GP, germination percentage.
Effect of papaya seed pre-sowing treatments on seed GP. Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. GP, germination percentage.

Figure 2.

Effect of papaya seed pre-sowing treatments on germination rate index (A), CV (B) and SVI (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. CV, coefficient of velocity; GRI, germination rate index; SVI, seedling vigour index.
Effect of papaya seed pre-sowing treatments on germination rate index (A), CV (B) and SVI (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. CV, coefficient of velocity; GRI, germination rate index; SVI, seedling vigour index.

Figure 3.

Effect of papaya seed pre-sowing treatments on seed antioxidant activity (DPPH%) (A), total phenolic content (mg · g−1 DW) (B) and total flavonoid content (mg · g−1 DW) (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. DPPH, 2,2-diphenyl-1-picrylhydrazyl.
Effect of papaya seed pre-sowing treatments on seed antioxidant activity (DPPH%) (A), total phenolic content (mg · g−1 DW) (B) and total flavonoid content (mg · g−1 DW) (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. DPPH, 2,2-diphenyl-1-picrylhydrazyl.

Figure 4.

Effect of papaya seed pre-sowing treatments on α-amylase (μmol · min−1 · mL−1) (A) and RS (mg · g−1 DW) (B). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. RS, Reducing sugar.
Effect of papaya seed pre-sowing treatments on α-amylase (μmol · min−1 · mL−1) (A) and RS (mg · g−1 DW) (B). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. RS, Reducing sugar.

Figure 5.

Effect of papaya seed pre-sowing treatments on proline content (mg · g−1 DW) (A) and total phosphorus (%) (B). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan.
Effect of papaya seed pre-sowing treatments on proline content (mg · g−1 DW) (A) and total phosphorus (%) (B). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan.

Figure 6.

Effect of papaya seed pre-sowing treatments on seedling height (cm) (A), number of roots (B) and root length (cm) (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan.
Effect of papaya seed pre-sowing treatments on seedling height (cm) (A), number of roots (B) and root length (cm) (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan.

Figure 7.

Effect of papaya seed pre-sowing treatments on stem diameter (A), number of leaves (B) and leaf area (cm2) (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan.
Effect of papaya seed pre-sowing treatments on stem diameter (A), number of leaves (B) and leaf area (cm2) (C). Data are means of two seasons (2022/2023 and 2023/2024) and three replicates (n = 3) each season. The mean values ± SE of each parameter followed by different alphabetical letters are significantly different according to Duncan’s multiple range test at p ≤ 0.05. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan.

Figure 8.

Electrophotograph generated by SDS-PAGE analysis of protein-banding pattern of untreated and submerged papaya seeds in magnetic water (2000 Gauss), M.W; chitosan (1.5%), CH and gibberellic acid (1000 ppm), GA3 separately or in combination. M, marker; Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan; K. Da, Kilo Dalton; SDS-PAGE, sodium dodecyl sulphate polyacrylamide gel electrophoresis.
Electrophotograph generated by SDS-PAGE analysis of protein-banding pattern of untreated and submerged papaya seeds in magnetic water (2000 Gauss), M.W; chitosan (1.5%), CH and gibberellic acid (1000 ppm), GA3 separately or in combination. M, marker; Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan; K. Da, Kilo Dalton; SDS-PAGE, sodium dodecyl sulphate polyacrylamide gel electrophoresis.

Figure 9.

Dendrogram derived from separate evaluation of SDS-PAGE protein profiles of eight treated and untreated papaya seeds constructed data using UPGMA and similarity matrices computed according to Dice coefficient. T1 (C), Control (dry seeds); T2, tap water (T.W); T3, magnetic water (M.W, 2000 Gauss); T4, chitosan (CH, 1.5%); T5, gibberellic acid (GA3, 1000 ppm); T6, M.W (2000 Gauss) + GA3 (1000 ppm); T7, M.W (2000 Gauss) + CH (1.5%); T8, GA3 (1000 ppm) + CH (1.5%); T9, M.W (2000 Gauss) + GA3 (1000 ppm) + CH (1.5%). SDS-PAGE, sodium dodecyl sulphate polyacrylamide gel electrophoresis; UPGMA, unweighted pair-group method with arithmetic averages.
Dendrogram derived from separate evaluation of SDS-PAGE protein profiles of eight treated and untreated papaya seeds constructed data using UPGMA and similarity matrices computed according to Dice coefficient. T1 (C), Control (dry seeds); T2, tap water (T.W); T3, magnetic water (M.W, 2000 Gauss); T4, chitosan (CH, 1.5%); T5, gibberellic acid (GA3, 1000 ppm); T6, M.W (2000 Gauss) + GA3 (1000 ppm); T7, M.W (2000 Gauss) + CH (1.5%); T8, GA3 (1000 ppm) + CH (1.5%); T9, M.W (2000 Gauss) + GA3 (1000 ppm) + CH (1.5%). SDS-PAGE, sodium dodecyl sulphate polyacrylamide gel electrophoresis; UPGMA, unweighted pair-group method with arithmetic averages.

Figure 10.

GTS% based on changes in SDS-PAGE protein electrophoretic banding pattern of untreated and pre-sowed papaya seeds in magnetic water (2000 Gauss), M.W; chitosan (1.5%), CH and gibberellic acid (1000 ppm), GA3 separately or in combination. Vertical bars represent GTS% ± standard error. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. GTS%, genome template stability percentage. SDS-PAGE, sodium dodecyl sulphate polyacrylamide gel electrophoresis.
GTS% based on changes in SDS-PAGE protein electrophoretic banding pattern of untreated and pre-sowed papaya seeds in magnetic water (2000 Gauss), M.W; chitosan (1.5%), CH and gibberellic acid (1000 ppm), GA3 separately or in combination. Vertical bars represent GTS% ± standard error. Control, dry seeds; T.W, tap water; M.W, magnetic water (2000 Gauss); CH, chitosan (1.5%); GA3, gibberellic acid (1000 ppm); M.W + GA3, magnetic water + gibberellic acid; M.W + CH, magnetic water + chitosan; GA3 + CH, gibberellic acid + chitosan; M.W + GA3 + CH, magnetic water + gibberellic acid + chitosan. GTS%, genome template stability percentage. SDS-PAGE, sodium dodecyl sulphate polyacrylamide gel electrophoresis.

Formula used to calculate papaya seed germination parameters, according to Taghizadeh and Solgi (2017)_

GPGP=NgNt×100{\rm{GP}} = {{{\rm{Ng}}} \over {{\rm{Nt}}}} \times 100Ng = germinated seedsNt = total sown seeds
GRIGRI=G11+G22++Gii{\rm{GRI}} = {{G1} \over 1} + {{G2} \over 2} + \ldots + {{Gi} \over i}Gi = the germination on each day after number of days from sowingi = alternate day after sowing
CVCV=ΣNiΣNi×Ti×100{\rm{CV}} = {{\Sigma Ni} \over {\Sigma Ni \times Ti}} \times 100Ni = the number of germinated seedsTi = the last day of germination
SVISVI = (PL + RL) × GPPL = shoot lengthRL = root lengthGP = germination percentage

SDS-PAGE analysis of protein-banding patterns of untreated and submerged papaya seeds in tap water; T_W, magnetic water (2000 Gauss); M_W, chitosan (1_5%); CH, gibberellic acid (1000 ppm); GA3 solutions separately or in combination; %P, % polymorphism_

No.Molecular weight (kDa)ControlTreatmentsBand type
T.WM.WCHGA3MW + ga3GA3 + CHMW + CHMW + GA3 + CH
1200.20110000000P
2178.12101111111P[NUM-2]
3171.50000000001P[PUM-9]
4151.72111111111M80
5102.25000000001P[PUM-9]
665.42010001000P
756.02010001110P
850.85000010000P[PUM-5]
946.92000010000P[PUM-5]
1043.73100100001P
1138.87001111000P
1236.64011110100P
1335.60100001111P
1433.42001000000P[PUM-3]
1527.73000101000P
1626.87100000000P[PUM-C]
1725.20100000000P[PUM-C]
1824.13111110111P[NUM-6]
1923.22000001001P
2019.46000010000P[PUM-5]
2118.82110000000P
2218.19000001010P
2315.23100001001P
2414.57111111101P[NUM-8]
2514.16000000010P[PUM-8]
2613.65010000010P
2711.54111110111P[NUM-6]
2810.93000100000P[PUM-4]
2910.58000100000P[PUM-4]
3010.46111111111M
3110.35100000000P[PUM-C]
3210.24000000001P[PUM-9]
Total bands of each sample 1411912111291013Gel polymorphism % = 93.75%
P% in each sample 85.7181.8177.7883.3381.8283.3377.7880.0084.62

Number of DS and S bands recorded by separate evaluation of protein profiles of treated and untreated papaya seeds and their genetic similarity indices as computed according to Dice coefficient_

TreatmentsControlT.WM.WCHGA3M.W + GA3GA3 + CHM.W + CHM.W + GA3 + CH
Control100%0DS,32S80%11DS,21S80%11DS,21S77%12DS,20S75%13DS,19S72%14DS,18S84%9DS,23S77%12DS,20S849DS,23S
T.W 100%0DS,32S868DS,24S79%11DS,21S81%10DS,22S75%13DS,19S90%26DS,6S84%9DS,23S75%13DS,19S
M.W 1000DS,32S92%5DS,27S93%4DS,22S80%11DS,21S93%4DS,28S84%9DS,23S82%10DS,22S
CH 100%0DS,32S88%7DS,25S77%12DS,20S88%7DS,25S77%12DS,20S79%11DS,21S
GA3 100%0DS,32S75%13DS,19S90%6DS,26S79%11DS,21S77%12DS,20S
M.W + GA3 100%0DS,32S84%9DS,23S86%8DS,24S79%11DS,21S
GA3 + CH 100%0DS,32S92%5DS,27S86%8DS,24S
M.W + CH 100%0DS,32S80%11DS,21S
M.W + GA3 + CH 100%0DS,32S

GTS% based on changes in SDS-PAGE protein electrophoretic banding pattern of untreated and submerged papaya seeds in magnetic water (2000 Gauss), M_W; chitosan (1_5%), CH; gibberellic acid (1000 ppm), GA3; separately or in combination_ a, number of newly developed bands; b, number of diminished bands; a + b, polymorphic bands

Control total bands Treatments
T.WM.WCHGA3M.W + GA3M.W + CHGA3 + CHM.W + GA3 + CH
abababababababab
14 4738575868274845
a + b 11 11 12 13 14 9 12 9
GTS%10021.43 21.43 14.29 7.14 0.00 35.71 14.29 35.71

Genomic-DNA template stability (GTS) and correlated % growth parameters and phytochemical changes among untreated and submerged papaya seeds_

GTS-Growth parameters
GTS%Seedling heightSVIStem diameterNumber of leavesRoot lengthLeaf areaRoot number% GerminationCV
GTS%1.00–0.17–0.09–0.16–0.13–0.10–0.090.220.10–0.05
GTS-Phytochemicals
GTSDPPH α-AmylaseProlineRSsTotal phenolicsFlavonoidsPhosphorous
GTS%1.000.29 –0.150.29–0.110.220.100.38*
DOI: https://doi.org/10.2478/fhort-2024-0036 | Journal eISSN: 2083-5965 | Journal ISSN: 0867-1761
Language: English
Page range: 533 - 558
Submitted on: Nov 21, 2024
Accepted on: Jan 21, 2025
Published on: Mar 27, 2025
Published by: Polish Society for Horticultural Sciences (PSHS)
In partnership with: Paradigm Publishing Services
Publication frequency: 2 issues per year

© 2025 M. S. Aboryia, Aml M. Abo AL-Saoud, El-Sayed A. El-Boraie, Hany G. Abd El-Gawad, Shaimaa Ismail, Mohamed Alkafafy, Bandar S. Aljuaid, Lina M. Abu-Ziada, published by Polish Society for Horticultural Sciences (PSHS)
This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 3.0 License.