Scientia Horticulturae 129 (2011) 353–360

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Physiological response of onion plants to foliar application of putrescine

and glutamine
A.A. Amin a , Fatma A.E. Gharib b , M. El-Awadi a , El-Sherbeny M. Rashad a,∗
a
Botany Department, National Research Centre, El-Tahrir St., El-Dokki, Cairo, Egypt
b
Botany Department, Faculty of Science, Helwan University, Cairo, Egypt

a r t i c l e i n f o a b s t r a c t

Article history: Two field experiments were carried out in Egypt during two successive seasons (2007/2008 and
Received 6 September 2010 2008/2009). This study aimed to investigate the response of growth, yield quality and some metabolic
Received in revised form 19 March 2011 constituents of onion (Allium cepa L. cv. ‘Giza 20’) to foliar application of putrescine (Put; 25, 50 and
Accepted 31 March 2011
100 mg L−1 ) and glutamine (Glut; 50, 100 and 200 mg L−1 ), the former a diamine and the latter an amino
acid, either alone, or in combination. Foliar application of Put and Glut, either alone or in combina-
Keywords:
tion, significantly increased plant height, number of leaves, fresh weight of leaves/plant, fresh and dry
Glutamine
weight/plant, leaf area, leaf area/plant, bulb length, bulb diameter and weight, as well as yield of onion
Growth
Onion plants
and quality of bulbs. Total soluble sugars, sulfur compounds, total soluble phenols, total free amino acids
Photosynthetic pigments and total photosynthetic pigment content in leaves were increased by increasing Put and/or Glut con-
Polyamines centrations up to 100 and 200 mg L−1 , respectively. Generally, foliar application of Put at 100 mg L−1 and
Putrescine Glut at 200 mg L−1 singly, or combined, effectively increased bulb yield and quality. In conclusion, the
Yield yield-contributing characters and quality of onion could be improved by application of Put and/or Glut.
© 2011 Elsevier B.V. All rights reserved.

1. Introduction
Onion (Allium cepa L.) is one of the most important vegetable
crops in the world. It is grown in Egypt mainly for local con-
sumption and export. Onion plants possess anti-bacterial and
anti-fungal activities, and contain powerful antioxidants (BHA;
butylated hydroxyanisole and BHT, butylated hydroxytoluene), sul-
fur and numerous other phenolic compounds (Singh et al., 2003;
Benkeblia, 2005). In onion bulbs ␥-glutamylpeptide and alliin con-
tents may account for up to 80% of the organic sulfur fraction. Alliins
and ␥-glutamylpeptides have therapeutic utility and might have
potential value as phytopharmaceutics. The alliins and their break-
down products (e.g. allicin) are the flavor precursors for the odor
and taste of Allium species (Saito et al., 2005).
Plant growth regulators (PGRs) regulate growth and devel-
opment. Polyamines (PAs) and amino acids are among the
“nontraditional” PGRs involved in metabolic processes. PAs are
polycationic compounds of low molecular weight that are present
in all living organisms, a category of PGRs involved in a large
spectrum of physiological processes, such as embryogenesis, cell
division, morphogenesis, floral initiation and development, and
stress (Paschalidis and Roubelakis-Angelakis, 2005; Liu et al., 2006;
Pang et al., 2007). PAs effectively retard senescence by retarding the
∗ Corresponding author.
E-mail address: sherbeny rashad56@yahoo.com (E.-S.M. Rashad).
loss of chlorophyll (Chl) (Couée et al., 2004; Tang et al., 2004). The
application of putrescine (Put) (up to 5 mM), a diamine without any
–COOH group, significantly increased the content of photosynthetic
pigments (Chl a, Chl b and total carotenoids) in leaves of wheat
(Triticum vulgare) plants (El-Bassiouny et al., 2008). PAs have been
shown to be an integral part of a plant’s response to stress (Alcázar
et al., 2006).
On the other hand, amino acids are the building blocks of
proteins and serve in a variety of important pathways. They
are important in many biological molecules, such as forming
parts of coenzymes, or as precursors for the biosynthesis of
molecules such as glutamine (Glut) and ornithine, which are pre-
cursors for nucleotides and PAs, respectively (Alcázar et al., 2010).
Amino acids are used for a variety of applications in industry
as an additive to animal feed, synthesis of drugs and cosmet-
ics (Leuchtenberger et al., 2005). The food technology is also
a major consumer of amino acids, particularly glutamic acid, a
common flavor enhancer (Garattini, 2000). In addition, amino
acids play an important role in plant “stress” resistance; glu-
tamate, cysteine and glycine are involved in biosynthesis of
glutathione (Meister, 1983). Glutathione plays a key role in detox-
ification of certain xenobiotics in higher plants (Rossini et al.,
1996), serves as a precursor of phytochelatins, which play an
important role in the detoxification of certain heavy metals in
plants (Rauser, 1995) and is involved in quenching free radicals
formed in plants under stress (Smirnoff, 1995). Put at 50 mg/l and
Glut at 200 mg/l were highly efficient in promoting growth and

0304-4238/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.scienta.2011.03.052
354 A.A. Amin et al. / Scientia Horticulturae 129 (2011) 353–360
yield parameters of faba bean (Vicia faba) plants (Rashad et al.,
2003).
The uses of PAs and amino acids to improve growth and yield
of various crops have yielded very encouraging results with some
plants: the addition of Put or arginine significantly increased fresh
and dry weights (FW and DW) of bean plants (Nassar et al., 2003).
Put at 10−5 M increased grain yield, biological yield and seed weight
index of wheat plants (Gupta et al., 2003). Exogenous application
of Put and Spd (spermidine) increased endogenous PGRs, partic-
ularly growth promoters (indole-3-acetic acid, gibberellins and
cytokinins) and decreased inhibitors (abscisic acid) in pea (Pisum
sativum) and wheat (T. vulgare) (El-Bassiouny, 2004 using 10−4 M
Put; Iqbal et al., 2006). Moreover, phenylalanine and ornithine (50
and 100 mg/l, respectively) significantly increased the FW and DW
of lemon grass (Cymbopogon nardus) (Gamal El-Din et al., 1997).
A high percentage of somatic embryogenesis and shoot regener-
ation of sugarcane (Saccharum officinarum) plants using glycine
(0.75 mM), arginine (0.5 mM) and cysteine (0.25 mM) was achieved
by Asad et al. (2009). Exogenously applied Put at 1.5 mM, Spd at
1.5 mM, 1.5 mM Put + 1.5 mM Spd, or 1.5 mM Put + 1.5 mM Spm
(spermine) significantly improved the growth rate of callus cul-
tures, shoot formation, and rooting of adventitious shoots (Tang
et al., 2004).
Considering the positive benefit that Put and Glut have had on
productivity of several crop plants, we decided to ascertain whether
these two compounds would also be able to increase morpho-
logical parameters of onion plants. By spraying onion plants with
different concentrations of Put and Glut, individually or in combi-
nation, the effect on some morphological criteria, yield and quality
of bulbs, as well as on some metabolic constituents of the plants
were assessed.
2. Materials and methods
Two field experiments were carried out at the Experimental
Station of the National Research Center in Nobaria region, Behira
Governorate, Egypt, during two successive seasons (2007/2008 and
2008/2009) to study the effect of foliar application of different con-
centrations of Put and Glut as well as their combination on growth,
photosynthetic pigments/leaves, yield and its quality and some bio-
chemical constituents of onion plants. Seeds of onion (A. cepa L. cv.
‘Giza-20’), recommended by the Egyptian Ministry of Agriculture
for sandy soil as occurs in the Nobaria region, were cultivated in a
nursery on the 21st November in both seasons (2006 and 2007).
The experimental design was a completely randomized block
design with four replications and 10 treatments. The plot area was
10.5 m2 (3.0 m × 3.5 m) and consisted of five rows 60 cm apart. After
70 days from sowing, seedlings were transplanted to an experimen-
tal field with a 10 cm inter-seedling spacing.
In both seasons, a foliar spray (pH 5.8) was applied twice to onion
plants during vegetative growth (at 30 and 50 days after transplant-
ing (DAT)), with Put at 25, 50 and 100 mg L−1 and/or Glut at 50, 100
and 200 mg L−1 (i.e. singly and in combination, in all possible per-
mutations). Solutions were sprayed evenly over the entire surface
of the plant (approx. 1 L plant−1 ), including the adaxial and abaxial
surfaces of leaves. No additives or surfactants such as Tween were
added. The interaction of different concentrations of both com-
pounds was also assessed, in addition to untreated plants (control),
which were sprayed only with tap water.
The plant growth characters that were measured at 90 DAT (i.e.
green yield) were: plant height, number of leaves/plant, FW of
leaves/plant, FW and DW/plant, and leaf area following the sug-
gestions of Bremner and Taha (1966), leaf area/plant, bulb length,
bulb diameter, FW and DW. Photosynthetic pigment (Chl a and b,
carotenoids as well as total pigments) content of fresh leaves was
also determined (Saric et al., 1967) at three stages of onion growth
(on 60, 75 and 90 DAT).
At the time of harvest (120 DAT) when 80% of the plant tops
had collapsed, the mean values of yield and its related parameters,
i.e. DW percentage/100 g DW, bulb length, bulb diameter, bulb FW
and DW and bulb yield (ton/1 fed [1 ha = 2.5 feddan (fed)]) were
determined. Plant samples were dried in an electric oven with a fan
at 70 ◦ C for 48 h until constant DW was achieved. Representative
dry samples of bulbs were taken from each treatment to determine
total soluble sugars using the phenol sulphuric method (Dubois
et al., 1956). Sulfur compounds were determined according to Singh
et al. (2003) and total soluble phenols according to A.O.A.C. (1990).
The method of Rosein (1957) was used to estimate total free amino
acids.
Data for both growing seasons was carried out according to
Snedecor and Cochran (1990). Data was combined since CV% was
<5%. Treatment means were compared using Fisher’s least signifi-
cant difference (LSD) at P = 0.05.
3. Results
3.1. Growth parameters
Data presented in Tables 1 and 2 shows that foliar applica-
tion of Put either separately at 25, 50 and 100 mg L−1 and Glut
at 50, 100 and 200 mg L−1 or their combination, promoted almost
all growth criteria compared to corresponding untreated plants. In
all cases, the increments in growth parameters were often highly
significant in comparison with untreated control plants. The most
effective treatments on growth parameters were the combination
of Put at 100 mg L−1 + Glut at 200 mg L−1 followed by Put alone at
100 mg L−1 .
When considered as a single factor, foliar application of Put
up to 100 mg L−1 significantly increased plant height, number of
leaves/plant, FW of leaves/plant, FW and DW/plant, leaf area, leaf
area/plant, bulb length, diameter and FW and DW compared to
untreated control plants.
Furthermore, when considering Glut as a single factor, it was less
effective than Put in increasing vegetative growth of onion plants
at the three different stages of growth (Table 1). The increment in
growth characters (i.e. plant height, number of leaves/plant, FW
of leaves/plant, fresh and DW/plant, leaf area, leaf area/plant, bulb
length, diameter and FW and DW was maximum with 200 mg L−1
Glut compared to control plants.
Regarding the combined effect of Put and Glut on vegetative
growth, similar significant increases were obtained in the same
growth parameters using different concentrations of Put and Glut
compared to control plants. Put at 100 mg L−1 + Glut at 200 mg L−1
enhanced growth parameters the most (Table 2).
3.2. Yield and quality of bulbs
Data presented in Tables 3 and 4 show that application of Put
increased onion yield and quality of bulbs more than control plants.
The most promising results, obtained with 100 mg L−1 Put, were for
bulb yield (15.86 ton/fed) and bulb quality (percent DW (16.87%),
bulb length (6.97 cm), bulb diameter (5.94 cm) and bulb weight
(169.93 g).
Glut alone up to 200 mg L−1 significantly increased onion yield
(ton/1 fed) and the quality of bulbs (i.e. dry weigh percentage, bulb
length, bulb diameter and FW and DW) compared to control plants
(Table 3).
Furthermore, onion yield was far more sensitive to the inter-
action between Put and Glut. The highest increase in onion yield
(15.86, 10.50 ton/fed) and bulb quality (i.e. %DW (16.92, 13.92%),
Table 1
Effect of putrescine and glutamine on growth characters of onion after 90 days from transplanting (combined analysis of two seasons).

Treatments (mg L−1 ) Growth parameters

Plant height Number of Leaves weight Fresh weight Dry weight (g) Leaf area Leaves Bulb length Bulb diameter Bulb weight
(cm) leaves (g) plant−1 (g) plant−1 plant−1 (cm2 ) area/plant(cm2 ) (cm) (cm) (g)

Putrescine Control 64.89 8.48 47.55 139.90 22.09 96.85 899.08 3.51 3.11 82.65
25 69.35 9.43 59.01 157.08 26.01 101.49 997.84 4.62 3.89 91.39
50 73.20 10.67 67.87 167.90 28.56 105.59 1130.06 4.93 4.76 97.83
100 76.05 10.89 69.35 170.95 29.41 110.44 1187.74 5.84 4.98 100.30
L.S.D. at 5% 4.06 0.88 10.46 13.09 2.32 4.01 123.09 0.29 0.16 2.33
Glutamine Control 65.01 8.31 46.67 138.88 21.79 94.98 796.62 3.31 3.09 80.69
25 67.90 9.01 57.91 155.72 25.73 99.55 905.15 4.37 3.54 88.40
50 71.82 9.89 64.15 165.09 26.98 102.39 1004.05 4.63 4.28 92.11

A.A. Amin et al. / Scientia Horticulturae 129 (2011) 353–360

100 74.10 10.32 67.06 168.87 28.01 106.62 1147.47 4.54 4.39 95.38
L.S.D. at 5% 2.56 0.67 8.24 11.84 1.12 3.55 96.21 0.24 0.10 1.86

Table 2
Effect of interaction between putrescine and glutamine on growth characters of onion after 90 days from transplanting (combined analysis of two seasons).

Treatments (mg L−1 ) Growth parameters

Putrescine Glutamine Plant height Number of Leaves weight Fresh weight Dry weight (g) Leaf area Leaves Bulb length Bulb diameter Bulb weight
(cm) leaves (g) plant−1 (g) plant−1 plant−1 (cm2 ) area/plant(cm2 ) (cm) (cm) (g)

0.0 0.0 60.11 7.71 53.21 146.40 21.49 94.18 726.13 3.51 2.91 70.78
50 64.26 8.42 60.76 152.92 24.56 97.27 819.01 4.62 3.31 81.50
100 67.34 8.69 62.86 159.51 27.80 100.36 827.13 4.73 3.42 84.21
200 69.49 8.94 64.90 167.79 30.11 106.45 951.66 5.48 3.73 86.89
25 0.0 65.58 8.59 61.43 155.11 25.90 96.54 829.28 4.90 3.64 88.53
50 69.78 8.96 62.71 166.29 29.41 99.63 892.68 4.71 3.75 91.21
100 71.78 9.37 64.59 169.38 32.01 103.72 971.86 5.57 4.66 103.39
200 75.86 9.89 68.11 172.47 36.40 109.81 1086.02 5.79 4.87 109.47
50 0.0 69.94 9.41 65.80 159.56 26.23 98.90 930.65 4.98 3.80 92.43
50 72.12 9.90 68.72 169.65 29.42 101.03 1001.97 5.64 3.92 96.30
100 75.20 10.11 70.62 172.67 34.61 106.52 1076.92 5.77 4.78 106.36
200 78.48 10.32 72.78 176.81 37.82 110.21 1137.36 5.96 4.91 111.29
100 0.0 71.56 9.49 66.56 167.09 28.19 100.30 951.85 5.70 4.52 105.23
50 75.64 10.14 69.89 170.01 33.28 104.49 1059.53 5.89 4.63 108.60
100 77.72 10.50 73.31 174.10 37.37 109.58 1150.59 6.29 4.82 112.01
200 79.80 11.06 76.20 178.23 39.46 112.67 1246.13 6.46 4.97 118.33
L.S.D. at 5% 2.01 0.22 1.77 3.09 2.39 3.42 54.18 0.13 0.10 2.29

355
 356
Table 3
Effect of putrescine and glutamine on yield, quality and chemical constituents of onion after 120 days from transplanting (combined analysis of two seasons).

Treatments (mg L−1 ) Yield and quality of onion Chemical constituents of onion

Bulb length Bulb diameter Bulb weight Dry weight Yield of onion Total soluble Free amino Total Sulfur
(cm) (cm) (g) (g/100 g. FW) (ton/fed) sugars % acid phenols compounds
mg/100(g) f.w

Putrescine Control 5.39 4.76 140.09 13.92 12.09 12.22 67.31 46.27 121.61
25 5.94 5.44 169.23 15.03 13.61 13.61 71.42 54.36 124.72
50 6.66 5.92 189.67 16.20 14.11 14.52 78.53 59.45 126.83
100 6.97 6.59 202.22 16.93 15.86 14.93 80.64 62.54 129.54
L.S.D. at 5% 0.29 0.41 11.50 0.56 1.52 0.35 2.61 3.01 2.69
Glutamine Control 5.33 4.70 139.79 13.87 11.96 11.96 66.57 44.63 120.52
25 5.76 4.89 157.83 14.68 13.06 12.41 69.68 50.72 123.61

A.A. Amin et al. / Scientia Horticulturae 129 (2011) 353–360

50 5.82 5.77 182.45 15.76 13.96 13.52 72.79 56.81 125.70
100 6.67 5.98 196.60 16.08 14.71 14.63 79.80 59.90 127.89
L.S.D. at 5% 0.25 0.17 9.34 0.31 0.76 0.26 2.11 2.89 1.96

Table 4
Effect of interaction between putrescine and glutamine on yield, quality and chemical constituents of onion after 120 days from transplanting (combined analysis of two seasons).

Treatments (mg L−1 ) Yield and quality of onion Chemical constituents of onion

Putrescine Glutamine Bulb length Bulb diameter Bulb weight Dry weight Yield of onion Total soluble Free amino Sulfur Total phenols
(cm) (cm) (g) (g/100 g. FW) (ton/fed) sugars % acids compounds (mg/100 g.DW)
(mg/100 g.DW)

0.0 0.0 4.31 3.52 119.79 11.83 10.50 11.26 60.21 119.81 43.28
50 4.62 3.71 121.76 12.05 11.66 12.34 64.34 122.62 47.67
100 5.43 4.62 132.19 13.21 12.78 12.66 68.43 125.39 49.56
200 5.64 4.79 139.67 13.79 12.89 12.80 70.65 127.48 51.75
25 0.0 4.58 3.90 125.48 12.39 11.62 12.60 63.46 121.55 45.61
50 5.60 4.68 130.90 12.96 12.86 13.43 69.77 124.46 49.42
100 5.79 4.87 136.71 13.51 13.72 13.72 71.89 126.76 53.69
200 6.43 4.90 141.62 14.01 13.93 14.33 74.56 128.58 56.78
50 0.0 5.76 4.59 138.29 13.66 12.64 13.74 67.40 123.69 48.46
50 5.87 4.84 145.80 14.40 13.85 13.85 70.63 126.40 53.73
100 5.92 5.56 156.29 15.46 13.97 14.60 75.82 129.21 57.82
200 6.60 5.79 163.31 16.29 14.67 15.39 78.86 131.26 59.90
100 0.0 6.49 4.77 142.69 14.09 13.56 14.80 69.70 125.30 50.68
50 6.59 5.60 158.70 15.77 14.48 14.96 74.88 127.49 54.76
100 7.36 5.79 163.42 16.34 15.63 15.01 79.59 130.58 59.84
200 7.47 6.94 169.93 16.97 15.92 15. 87 81.01 132.67 62.92
L.S.D. at 5% 0.16 0.14 4.31 0.52 0.12 0.09 1.49 1.54 1.29
 A.A. Amin et al. / Scientia Horticulturae 129 (2011) 353–360 357

bulb length (7.74, 5.39 cm), bulb diameter (6.59, 4.76 cm) and bulb
DW (169.93, 140.09 g) were obtained by foliar application with

pigments
100 mg L−1 Put + 200 mg L−1 Glut (Table 4) followed by 100 mg L−1

Total

1.25

1.49
1.58

1.34
1.43
1.52
1.40

0.09
1.20

0.06
Put (15.82 ton/fed) for all previous parameters (Table 3). In all cases,
the increments in onion yield and bulb quality were often highly
significant in comparison with untreated controls.

0.43
0.49
0.52
0.56

0.42
0.46

0.53
0.03

0.50

0.02
Car.
Effect of putrescine and glutamine on the photosynthetic pigments content (mg/g fresh weight) in the leaves of onion plants at 60, 75, and 90 days after transplanting (combined analysis of two seasons).
3.3. Photosynthetic pigments

Chl. a + b
At 90 days after transplanting
Data presented in Tables 5 and 6 show that Chl a and b,

0.82
0.91
0.97

0.78
0.88
0.93
0.99
1.02
0.06

0.04
carotenoids and total photosynthetic pigments in the leaves of
onion plants reached a maximum value at 75 DAT. Generally, foliar
application of either Put or Glut at any concentration or their

Chl. b

0.23
0.24
0.25
0.26

0.22
0.24
0.24
0.25
0.01

0.01
combination significantly increased the Chl a + b, carotenoids and
consequently the total pigment contents more than controls at
60, 75 and 90 DAT. The most effective concentration was 100 and

Chl. a
200 mg L−1 of either Put or Glut, respectively and their combina-

0.59
0.67
0.72
0.76

0.56
0.64
0.69
0.74
0.05

0.03
tion at the three stages of growth (i.e. 10, 25 and 40 days after the
second spray of Put or Glut).
Foliar spray of onion plants at all three Put concentrations (25,

pigments
50 and 100 mg/l) tested significantly increased photosynthetic pig-

Total

1.37

1.69
1.81
0.11
1.39
1.49

1.71
1.50

1.60

0.08
ments compared to untreated plants (Table 5). The highest recorded
value of Chl a + b, carotenoid and total pigments in the leaves of
onion plants was obtained with 100 mg L−1 Put at the three stages of

0.39
0.44
0.49
0.52

0.42
0.47
0.03
0.40

0.50
0.02
growth, although Glut up to 200 mg L−1 also significantly increased

Car.
these parameters.
Moreover, the interaction between Put and Glut significantly

Chl. a + b
At 75 days after transplanting
increased Chl a and b, carotenoids and consequently the total pig-

0.98

1.29

0.99

1.13
1.21
1.06
1.20

0.08

1.07

0.06
ment content in the leaves of onion plants more than controls at
all three stages of growth (Table 6), the most effective treatment
being Put at 100 mg L−1 + Glut at 200 mg L−1 .

Chl. b

0.26
0.27
0.31
0.33

0.26
0.29

0.32
0.02

0.30

0.01
3.4. Chemical constituents
Chl. a
Data presented in Tables 3 and 4 show that foliar application of

0.72
0.79
0.89
0.96

0.73
0.78
0.83
0.89
0.06

0.05
either Put or Glut at any concentration as well as their combination
significantly increased the total sugar content, free amino acids,
phenolic and sulfur compounds in the dry bulbs of onion com-
pigments

pared with their controls at 120 DAT, the harvest date. The most
effective treatment was 100 and 200 mg L−1 of either Put or Glut,
Total

1.32
1.45
1.59
1.69

1.32
1.42

1.58
0.08

1.50

0.05
respectively or in combination.
Photosynthetic pigments (mg/g fresh weight)

Furthermore, the results obtained indicate that the highest level

of total phenolic compounds (61.92, 59.90 and 62.92 mg/100 g DW)
0.37

0.46
0.49

0.36
0.39
0.43
0.47
0.40

0.02

0.01
Car.

and sulfur compounds (129.54, 127.89 and 132.67 mg/100 g DW)

was obtained for either 100 mg L−1 Put, 200 mg L−1 Glut or their
Chl. a + b

combination, respectively compared with 43.28, 119.81 mg/100 g

At 60 days after transplanting

DW for controls.
0.95

1.13

0.96

1.11
1.05

1.20
0.06

1.03
1.07

0.04

4. Discussion
Chl. b

0.25
0.27
0.29

0.25
0.27
0.28
0.29
0.30
0.01

0.01

The present study indicates that the application of Put up to

100 mg L−1 and Glut up to 200 mg L−1 , individually or in combi-
nation, greatly promoted the vegetative growth and dry matter
Chl. a

0.78
0.84

0.71
0.76
0.79
0.82
0.70

0.90
0.05

0.03

production of onion by enhancing cell division and chlorophyll

accumulation. In other crops, Put also stimu1ated growth by
increasing the amount of endogenous promoters (auxin, gib-
Control

Control

berellins and cytokinins) accompanied by a decrease in the content

100

100
25

25
50

50

and activity of inhibitors (ABA) (El-Bassiouny and Bekheta, 2005;

El-Bassiouny et al., 2008). Put has shown a much better ability than
Glut (200 mg L−1 ) to promote growth (Nassar et al., 2003; Amin
Treatments (mg L−1 )

et al., 2008). However, the application of Put has only been reported
in a few cases: jimson weed (Datura stramonium) callus and lemon-
L.S.D. at 5%

L.S.D. at 5%
Glutamine

grass (C. nardus) plants (Moursy et al., 1988; Gamal El-Din et al.,
Pigments

1997). Intuitively, the combination of Put and Glut more effectively

Table 5

increased vegetative growth and dry matter production of onion

plants than single treatments.
358 A.A. Amin et al. / Scientia Horticulturae 129 (2011) 353–360

Effect of interaction between putrescine and glutamine on the photosynthetic pigments content (mg/g fresh weight) in the leaves of onion plants at 60, 75 and 90 days after transplanting (combined analysis of two seasons). Photosynthetic pigments of onion leaves were significantly
enhanced by the application of Put or Glut and/or their combina-

pigments
tion. Put and Glut might retard Chl destruction and/or increase their
Total

1.33

1.45

1.39
1.44
1.51
1.58
1.44
1.48
1.54

1.51
1.59
1.65
1.69
1.40

1.50

1.60

0.05
biosynthesis or stabilize the thylakoid membrane; PAs may retard
senescence by altering the stability and permeability of mem-
0.44 branes by protecting and preventing chloroplasts from senescing
0.47
0.49
0.52
0.47
0.49
0.52
0.56
0.49
0.51
0.53
0.56
0.52
0.56
0.59
0.60
0.03
Car.

and therefore retarding Chl loss (Gonzalez-Aguilar et al., 1997).

Diamine aminotransferase transfers the amino group of Put to ␣-
oxoglutaric acid, the precursor of Chl (Askar and Treptow, 1986),
Chl. a + b

the direct effect on Chl is not clear, although PAs have been show
0.93
0.96
0.98
0.92
0.95
0.99
1.02
0.95
0.97
1.01
1.04
0.99
1.03
1.06
1.09
0.02
90 days after transplanting

089

to affect Chl stabilization and antisenescence (Besford et al., 1993;

Legocka and Zajchert, 1999; Teixeira da Silva, 2006; Ioannidis
et al., 2007). Put increased endogenous cytokinins, which stim-
Chl. b

ulate Chl biosynthesis and chloroplast differentiation in wheat

0.23
0.24
0.25
0.25
0.24
0.25
0.26
0.26
0.25
0.25
0.26
0.27
0.25
0.26
0.27
0.28
0.01
(T. vulgare) (Xie et al., 2004; El-Bassiouny et al., 2008). Promo-
tive effects of PAs on photosynthetic pigments was observed for
mulberry (Morus alba L.) (Das et al., 2002), cucumber (Cucumber
Chl. a

0.66
0.69
0.71
0.73
0.68

0.73
0.76

0.72
0.75
0.77
0.74
0.77
0.79
0.81
0.70

0.70

0.02
sativus L.) (He et al., 2002) and common bean (Phaseolus vulgare)
(Nassar et al., 2003). Amino acid and Put treatments concomi-
tantly with increasing Mg levels (a structural component of Chl)
pigments

retarded senescence (Kao, 1994; El-Bassiouny et al., 2008). There-

fore, the combination of Put at 100 mg L−1 + Glut at 200 mg L−1 was
Total

1.37
1.45
1.53
1.59
1.48
1.57
1.64

1.54
1.63
1.74
1.78
1.63
1.72
1.79
1.87
1.70

0.05

the best treatment to increase photosynthetic pigments of onion

leaves.
Onion yield and the quality of bulbs was significantly enhanced
0.39
0.42
0.46
0.49
0.43
0.48
0.52
0.54
0.46
0.49
0.54
0.56
0.49
0.52
0.55
0.59
0.03
Car.

by Put, especially at higher concentrations. Put may increase

the existing PA pool and subsequently retard senescence. Rice
grain filling, plumpness and 1000-seed weight were positively
Chl. a + b

correlated with PA content (Yang et al., 1996). On the other

1.12
1.16

1.14

1.22
1.14

1.24
1.28
0.98
1.03
1.07
1.10
1.05
1.09

1.08

1.20

1.20

0.02
75 days after transplanting

hand, the application of amino acids has only been reported to

increase essential oil yield of lemon grass (C. nardus) (Gamal El-
Din et al., 1997). The Put + Glut combination was more effective
Chl. b

0.26
0.27
0.28
0.29
0.27
0.29

0.31
0.28

0.31
0.32

0.31
0.32
0.33
0.30

0.30

0.30

0.01

than Glut alone for increasing yield by enhancing photosynthetic

activity, accumulating dry matter and retarding senescence. Con-
sequently, there was increased translocation and accumulation of
Chl. a

certain metabolites in plant organs which affected yield and qual-

0.72
0.76
0.79
0.81
0.78

0.82
0.85

0.84
0.89

0.84
0.89
0.92
0.95
0.80

0.80

0.90

0.01

ity.
A foliar application of either Put or Glut and their combina-
tion increased the total sugar content, free amino acids, phenolic
pigments

and sulfur compounds in the dry bulbs of onion notably due to

the bioregulatory effect on enzymatic activity and translocation
Total

1.29
1.38
1.44

1.41
1.47
1.54
1.65
1.47
1.54
1.64
1.72
1.55
1.62
1.69
1.78
1.50

0.06

processes from leaves to bulbs, linking or converting to other

plant metabolites. PAs are linked with particular proteins (Serafini-
Fracassini and Del Duca, 2008). Covalently bound PA–protein
0.36
0.39

0.43
0.39
0.42
0.46
0.49
0.41
0.44
0.48
0.52
0.44
0.46
0.49
0.55
0.40

0.03
Car.

complexes have been reported in Helianthus tuberosus (artichoke)

tubers (Dinnella et al., 1992) and tobacco (Nicotiana spp.) (Sawhney
and Appeluthite, 1992). PAs are able to bind with pectin and
Chl. a + b

polysaccharides (D’oraci and Bagni, 1987) and increased ␣-amylase

1.16

1.16

1.11
1.16

1.23
0.93
0.99
1.04
1.07
1.02
1.05
1.08

1.06
1.10

1.20

1.20

0.03
60 days after transplanting

activity (Tipirdamaz et al., 1995). On the other hand, Spm and

Put significantly increased the total sugar content, total pheno-
lic compounds but decreased total free amino acids in chamomile
Chl. b

0.24
0.25
0.27
0.28
0.26
0.27
0.28

0.27
0.28

0.31
0.29

0.31
0.32
0.30

0.30

0.30

0.01

(Matricaria chamomilla) leaves (Abd El-Wahed and Gamal El-Din,

2004, 2005). The combination of Put and Glut had a high effect
on amino acid conversion to phenolic and sulfur-containing com-
Chl. a

pounds. Phenolic acids conjugate with PAs and modulate the

0.69
0.74
0.77
0.79
0.76
0.78

0.86
0.79
0.82
0.86
0.89
0.82
0.86
0.89
0.91
0.80

0.02

free levels of these plant growth substances. Thus, the effect

of L-␣ aminooxy-␤ phenylpropionic acid found on PAs (Martin-
Chl, chlorophyll; Car, carotene.

Tanguy, 1985) suggest that an interaction between phenolic acids

Glutamine

and PAs was involved in growth promotion by L-␣ aminooxy-␤

phenylpropionic acid (Mader and Hanke, 1997). Thus, combined
Treatments (mg L−1 )

100
200

100
200

100
200

100
200
0.0

0.0

0.0

0.0
50

50

50

50

treatments of Put at 100 mg L−1 and Glut at 200 mg L−1 was very
effective for enhancing total sugar content, total free amino acids,
L.S.D. at 5%
Putrescine

total phenolic and sulfur-containing compounds and consequently

may have directly increased the level of antioxidants, flavor pre-
Table 6

100

cursors for the odor and taste of onion bulb more than Put
0.0

25

50

or Glut used alone. ␥-Glutamyl peptides and alliins – the fla-

 A.A. Amin et al. / Scientia Horticulturae 129 (2011) 353–360
vor precursors for the typical odor and taste of onion bulbs –
may account for as much as 80% of the organic sulfur fraction
(Saito et al., 2005). Thus, the yield-contributing characters and
yield of onion could be manipulated by application of Put and/or
Glut.
In conclusion, Put and Glut can be easily applied to onion plants
in the field. Application of Put at 100 mg L−1 and Glut at 200 mg L−1 ,
singly or in combination, resulted in a significant increase in every
morphological attribute, including bulb fresh and dry weight, the
main consumer-related traits. The use of Put and Glut thus opens
up a new avenue for increasing yield and improving the quality of
an onion crop.
Acknowledgements
The authors thank Dr. Teixeira da Silva for substantial contri-
butions to interpretation of data, revising the article critically for
important intellectual content and for final approval of the ver-
sion, which had been previously and conditionally accepted to be
published.
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