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

, , , , , ,  und   
27. März 2025

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COVER HERUNTERLADEN

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)_

GP GP=NgNt×100{\rm{GP}} = {{{\rm{Ng}}} \over {{\rm{Nt}}}} \times 100 Ng = germinated seedsNt = total sown seeds
GRI GRI=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
CV CV=ΣNiΣNi×Ti×100{\rm{CV}} = {{\Sigma Ni} \over {\Sigma Ni \times Ti}} \times 100 Ni = the number of germinated seedsTi = the last day of germination
SVI SVI = (PL + RL) × GP PL = 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) Control Treatments Band type
T.W M.W CH GA3 MW + ga3 GA3 + CH MW + CH MW + GA3 + CH
1 200.20 1 1 0 0 0 0 0 0 0 P
2 178.12 1 0 1 1 1 1 1 1 1 P[NUM-2]
3 171.50 0 0 0 0 0 0 0 0 1 P[PUM-9]
4 151.72 1 1 1 1 1 1 1 1 1 M80
5 102.25 0 0 0 0 0 0 0 0 1 P[PUM-9]
6 65.42 0 1 0 0 0 1 0 0 0 P
7 56.02 0 1 0 0 0 1 1 1 0 P
8 50.85 0 0 0 0 1 0 0 0 0 P[PUM-5]
9 46.92 0 0 0 0 1 0 0 0 0 P[PUM-5]
10 43.73 1 0 0 1 0 0 0 0 1 P
11 38.87 0 0 1 1 1 1 0 0 0 P
12 36.64 0 1 1 1 1 0 1 0 0 P
13 35.60 1 0 0 0 0 1 1 1 1 P
14 33.42 0 0 1 0 0 0 0 0 0 P[PUM-3]
15 27.73 0 0 0 1 0 1 0 0 0 P
16 26.87 1 0 0 0 0 0 0 0 0 P[PUM-C]
17 25.20 1 0 0 0 0 0 0 0 0 P[PUM-C]
18 24.13 1 1 1 1 1 0 1 1 1 P[NUM-6]
19 23.22 0 0 0 0 0 1 0 0 1 P
20 19.46 0 0 0 0 1 0 0 0 0 P[PUM-5]
21 18.82 1 1 0 0 0 0 0 0 0 P
22 18.19 0 0 0 0 0 1 0 1 0 P
23 15.23 1 0 0 0 0 1 0 0 1 P
24 14.57 1 1 1 1 1 1 1 0 1 P[NUM-8]
25 14.16 0 0 0 0 0 0 0 1 0 P[PUM-8]
26 13.65 0 1 0 0 0 0 0 1 0 P
27 11.54 1 1 1 1 1 0 1 1 1 P[NUM-6]
28 10.93 0 0 0 1 0 0 0 0 0 P[PUM-4]
29 10.58 0 0 0 1 0 0 0 0 0 P[PUM-4]
30 10.46 1 1 1 1 1 1 1 1 1 M
31 10.35 1 0 0 0 0 0 0 0 0 P[PUM-C]
32 10.24 0 0 0 0 0 0 0 0 1 P[PUM-9]
Total bands of each sample 14 11 9 12 11 12 9 10 13 Gel polymorphism % = 93.75%
P% in each sample 85.71 81.81 77.78 83.33 81.82 83.33 77.78 80.00 84.62

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_

Treatments Control T.W M.W CH GA3 M.W + GA3 GA3 + CH M.W + CH M.W + GA3 + CH
Control 100%0DS,32S 80%11DS,21S 80%11DS,21S 77%12DS,20S 75%13DS,19S 72%14DS,18S 84%9DS,23S 77%12DS,20S 849DS,23S
T.W 100%0DS,32S 868DS,24S 79%11DS,21S 81%10DS,22S 75%13DS,19S 90%26DS,6S 84%9DS,23S 75%13DS,19S
M.W 1000DS,32S 92%5DS,27S 93%4DS,22S 80%11DS,21S 93%4DS,28S 84%9DS,23S 82%10DS,22S
CH 100%0DS,32S 88%7DS,25S 77%12DS,20S 88%7DS,25S 77%12DS,20S 79%11DS,21S
GA3 100%0DS,32S 75%13DS,19S 90%6DS,26S 79%11DS,21S 77%12DS,20S
M.W + GA3 100%0DS,32S 84%9DS,23S 86%8DS,24S 79%11DS,21S
GA3 + CH 100%0DS,32S 92%5DS,27S 86%8DS,24S
M.W + CH 100%0DS,32S 80%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.W M.W CH GA3 M.W + GA3 M.W + CH GA3 + CH M.W + GA3 + CH
a b a b a b a b a b a b a b a b
14 4 7 3 8 5 7 5 8 6 8 2 7 4 8 4 5
a + b 11 11 12 13 14 9 12 9
GTS% 100 21.43 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 height SVI Stem diameter Number of leaves Root length Leaf area Root number % Germination CV
GTS% 1.00 –0.17 –0.09 –0.16 –0.13 –0.10 –0.09 0.22 0.10 –0.05
GTS-Phytochemicals
GTS DPPH α-Amylase Proline RSs Total phenolics Flavonoids Phosphorous
GTS% 1.00 0.29 –0.15 0.29 –0.11 0.22 0.10 0.38*
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