Induced systemic resistance to Meloidogyne spp by <i>β</i>,amino butyric acid in tomato

doi:10.4236/as.2013.411081

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Vol.4, No.11, 608-613 (2013) Agricultural Sciences

http://dx.doi.org/10.4236/as.2013.411081

Induced systemic resistance to Meloidogyne spp by

β,amino butyric acid in tomato

Shamaael S. Mutar, Farkad A. Fattah*

Department of Plant Protection, College of Agriculture, University of Baghdad, Baghdad, Iraq;

*Corresponding Author: [email protected]

Received 17 September 2013; revised 16 October 2013; accepted 27 October 2013

tribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is prop-

erly cited.

ABSTRACT

β,amino butyric acid (BABA) induced resistance

against Meloigogyne spp in tomato. Signifi-

cantly (p = 0.05) less, 41.11 second stage juve-

niles (J2) enter the roots of treated than, 116.66

J2 in untreated control plants. Root fresh and

dry weight (Rfw, Rdw) were 2.87 and 0.12 g in

treated compared with 4.78 and 0.30 g in nema-

tode infected control plants respectively, 30 d

after nematode inoculation. Sfw and Sdw were

8.62 and 0.60 g compared with 3.94 and 0.22 g in

control plants. Foliage spray at 40 and 20 mM of

BABA was more effective than 10 and 5 mM

treatments. The former two concentrations re-

corded the lowest, average gall index, 2.11 com-

pared with 3.33, 4.11, and 5 for the latter two

concentrations and nematode infected control

respectively. Results also indicated that treat-

ments with BABA prior to nematode inoculation

were superior in inducing resistance to Meloi-

dogyne spp over treatment at the time or after

nematode inoculation.

Keywords: Induced Resistance; β,Amino Butyric

Acid; Meloigogyne spp; Tomato

1. INTRODUCTION

Tomato, Solanum lycopersicum L., formerly known as

Lycopersicon esculentum Mill., is the second most im-

portant vegetable crop after potato with annual world

production of about 152.9 million ton in 2009 [1]. An-

nual tomato production in Iraq was estimated at 830.000

tons in 2008 [2]. Tomato plants are subjected to infection

by many important plant pathogens including the root

knot nematodes, Meloidogyne spp which are considered

as the most important nematode species worldwide and

in Iraq. Many effective control measures were used to

manage these pathogens such as soil solarization chemi-

cal and biological control. Induced systemic resistance to

plant pathogen provides an ideal control measures

against plant pathogens [3]. Chemical, physical and bio-

logical inducers to control the root knot nematodes were

used [4]. In recent years, various chemical, physical and

biological agents were used to induce resistance to many

plant pathogens. β-1,3 amino butyric acid (BABA), Ben-

zol [1-3] thaiadiazole-7-carbothionic acid S-methyl ester

and salicylic acid are among the most widely studied

chemical inducers [5-7]. These chemicals activate resis-

tance against different plant pathogens like fungi, bacte-

ria, viruses and nematodes [8]. BABA was reported to

induce resistance against members of solanacea, cucur-

bitacea, legumenacea, cruciferea, and rosacea [9]. Jakob

et al. [10], reported that BABA induced resistance to dif-

ferent plant pathogens such as fungi, bacteria, viruses,

and plant parasitic nematodes. Only few studies on in-

duced resistance by BABA against plant nematodes are

reported. BABA was reported to reduce damage due to

root knot nematode on tomato [11] and M. javanica and

Rotylenchulus reniformis on pineapple [12] and induce

resistance against M. javanica in cucumber [4].

This study was undertaken to assess the ability of β,

aminobutyric acid to induce acquired systemic resistance

against the root knot nematodes, Meloidogyne spp in to-

mato plants.

2. MATERIALS AND METHODS

Experiments were performed in a greenhouse (27 ± 5

C) and tomato, Solanum lycopersicum L. cv. “Supper Re-

gina” highly susceptible to Meloidogyne spp was used.

Plants were grown and maintained in1kg plastic pots

throughout th e experiments.

2.1. Nematode Inoculation

Soil from cucumber grown plastic house heavily in-

S. S. Mutar, F. A. Fattah / Agricultural Sciences 4 (2013) 608-613 609

fested with Meloidogyne spp (predominantly M.javanica)

were collected and stored at 4 C in polyethylene pages

until use to inoculate tomato seedlings.The nematode

infested soil was mixed with peat moss in 1:1 ratio and

used as nematode inoculums.

2.2. Preparation β,Amino Butyric Acid

(BABA)

Stalk solution of BABA, Mwt = 103 (Sigma Aldrich,

St Louis, Missouri, USA) was prepared by dissolving

412 mg in 100 ml of distilled water.

2.3. Foliage Treatments with BABA

Four concentrations of BABA, 5, 10, 20, and 40 mM

were used as foliage application. The experiment was

consisted of 13 treatments (one nematode inoculated

control sprayed with distilled water and 12 combination

of the 4 concentrations of BABA 3-d before, during and

7-d after nematode inoculation). Plants were carefully

sprayed till complete wetness using 1L hand sprayer.

Each treatment was replicated 4 times and arranged in

randomized complete blocks in the greenhouse. Thirty

days after inoculation, plants were assessed for growth

and nematode infection.

2.4. Root Treatments with BABA

In this experiment, 2 mL of 1, 2.5 and 5 mM of BABA

were used as soil drenching added around the stem of

single tomato plants growing in n ematode infested so il in

1 kg plastic pots. Ten treatments (one nematode inocu-

lated control watered with 2 ml of distilled water and 9

combinations of the 3 concentration of BABA applied

3-days before, during and 7-days after nematode inocula-

tion ). Thirty days after inoculation, plants were assessed

for growth and nematode infection. Each treatment was

replicated 4 times and arranged in a randomized com-

plete block in the greenhouse. Thirty days after inocula-

tion, plants were assessed for growth and nematode in-

fection.

2.5. Effect of BABA on Meloidogyne spp

Rate of Gall Index

The rate of infestation of the root knot nematodes is

determined using a 5 level scales [13]: 1 = no galls on

roots, 2 = galls on 1% - 25% of the root, 3 = galls on

26% - 50% of the root, 4 = galls on 51% - 75% of the

root, and 5, galls on 76% - 100% of the root.

2.6. Effect of BABA on Fresh and Dry

Weight of Shoot and Root Systems

Plants were carefully uprooted and roots were washed

under tap water to remove adhering soil. To determine

shoot and root dry and wet weight, shoots and roots were

separately weights and dried at 70 C for 48 h or until

weight fixed.

2.7. Effect of BABA on Nematode

Penetration

To determine the effect BABA treatments on nematode

penetration of tomato roots, roots were stained with acid

fuchsine [14], washed with water to remove excess stain

and 1 g of each root was individual examined under a

compound microscope to count nematodes inside the

roots.

3. STATISTICAL ANALYSIS

The data were subjected to analysis of variance and

means were separated by the least significant method at

(p = 0.05) using SAS, 200 4 [1 5].

4. RESULTS

4.1. Foliage Application

4.1.1. Effect of Foliage Treatment of BABA on

Meloidogyne spp Rate of Gall Index

BABA treatment induced resistant to Meloidogyne spp

in tomato plants (Ta bl e 1). The treatments significantly

(p = 0.05) reduced the rates of gall index of treated plants

compared with nematode infected control plants. While

40, 20, 10, and 5 nM of BABA treated plants recorded

low average gall indices, 2.11, 2.11, 3.33, 4.11 respec-

tively. BABA untreated plants recorded the highest av-

erage gall index, 5, 30 days after nematode inoculation.

The former two concentrations render tomato plant sig-

Table 1. Effect of spraying BABA on tomato, Solanum ly-

copersicum L. shoot on rate of root galls index of Meloidogyne

spp infected plants 30 days after nematode inoculation.

Rate of root gall index

Nematode inocu l ation

BABA Conc.

(mM)

7 days beforeDuring 3 days after

Mean

40 2.33 2.33 1.66 2.11

20 2.66 2 1.66 2.11

10 4 3 3 3.33

5 4 4 4.33 4.11

Control 5 5 5 5

Mean 3.60 3.26 3.13

LSD (P = 0.05) = Conc. = 0.35*,Trea. = 0.27*, Inter = 0.60*. Each number is

a mean of t hree r epli cates an d two plan t each. *Indicate significant deferens.

Nematode inoculums (nematode infested soil + peat moss, 1:1) were added

when plants were 4 - 5 true leaves. Gall index was according to 1 - 5 level

scale: 1 = no galls on the roots, 2 = galls on 1% - 25% of the root, 3 = galls

on 26% - 50% of the root, 4 = galls on 51% - 75% of the root, and 5 = galls

on 76% - 100% of roots. BABA = β,amino butyric acid.

S. S. Mutar, F. A. Fattah / Agricultural Sciences 4 (2013) 608-613

610

nificantly more resistant to the nematode than the two

latter concentrations (Table 1). The 40 and 20 mM

scored significantly (p = 0.05) the lowest nematode gall

index, 1.66, when BABA treatments exceeded nematode

inoculation by 3 days compared with other test treat-

ments.

4.1.2. Effect of BABA on Fresh and Dry Weight

of Root Systems

Results in Tabl e 2 sho wed that spraying tomato p lants

with BABA significantly (p = 0.05) affected root fresh

weight (Rfw) and root dry weight (Rdw) of Meloidogyne

spp infected tomato plants. The 40 mM recorded the

lowest average root weights, 5.74 and 0.34 g compared

with other test treatments 30 days after nematode inocu-

lation. Treatment with BABA 3 days before nematode

inoculation produced the lowest Rfw and Rdw, 3.96 and

0.29 g respectively 30 days after nematode inoculation.

The 40 mM concentration also recorded the lowest root

weights, 2.35 and 0.21 g compared with the highest root

weight, 12.98 and 0.80 g when the treatment was per-

formed 7 days after nematode inoc ul a t i on.

4.1.3. Effect of BABA on Fresh and Dry Weight

of Shoot Systems

Spraying of BABA on foliage of tomato plants signifi-

cantly (p = 0.05) increased root fresh and dry weight

(Sfw and Sdw) of nematode infected tomato plants 30

days after inoculation (Table 3). The average w eights f or

plants treated with 40 and 20 mM of BABA r ecorded the

highest, 21.40 and 2.10 g Sfw and Sdw compared with

the lowest average weights of 15.05 and 0.97 g in nema-

tode infected controls. Treatment with BABA 3 days be-

fore nematode inoculation caused significantly the high-

est weight, 20.99 and 2.24 g compared with 14.24, 0.75,

19.31 and 1.35 g when nematode was inoculated during

and 7 days after BABA treatment respectively. Treatment

with 40 and 20 mM of BABA 3 days prior to nematode

inoculation caused higher Sfw and Sdw, 25.32, 3.25 and

22.22, 3.90 g compared with, 12.86 and 0.51 g for nema-

tode infected control plants respectively.

4.2. Soil Application

4.2.1. Effectof Soil Drenching with BABA on

Meloidogyne spp Rate of Gall Index

Soil drenching with BABA significantly (p = 0.05)

reduced the root ga ll ind ex of to mato plan ts infected with

Meloidogyne spp (Tab le 4). Gall index was decreased as

the test BABA concentration was increased. The 5 mM

concentration produced significantly the lowest average

gall index, 2.33 compared with, 2.77, 3.55 and 5 for 2.5,

1 mM and control 30 days after nematode inoculation r e-

spectively. The lowest average gall index, 2.91 was re-

corded when BABA was applied 3 days before nematode

Table 2. Effect of spraying BABA on tomato, Solanum ly-

copersicum L. shoot on root weights of Meloidogyne spp in-

fected plants 30 days after nematod e inocu lation.

Root weight of infected plants (g)

Nematode inocu l ation

7 days afterDuring 3 days before

BABA

Conc.

(mM)

DryFreshDryFresh Dry Fresh

Mean

Dry Mean

Fresh

40 0.212.350.375.97 0.45 8.90 0.345.74

20 0.242.720.405.98 0.47 10.09 0.376.26

10 0.283.910.506.74 0.51 10.16 0.436.94

5 0.305.890.728.92 0.50 11.44 0.518.75

Control0.404.940.528.79 0.80 12.98 0.588.90

Mean 0.293.960.507.28 0.54 10.71

LSD (P = 0.05) Dry weight: Conc. = 0.065*, Treat. = 0.050*, Inter. = 0.11*.

LSD (P = 0.05) Fresh weight: Conc. = 1.78*, Traet. = 1.37*, Inter. = 3.08*.

Each number is a mean of three replicates and two plant each. *Indicate

significant deferens. Nematode inoculums (nematode infested soil + peat

moss, 1:1) were added when plants were 4 - 5 true leaves. BABA = β,amino

butyric acid.

Table 3. Effect of spraying BABA on tomato, Solanum ly-

copersicum L. shoot on shoot weights of Meloidogyne spp in-

fected plants 30 days after nematod e inocu lation.

Shoot weight of infected plants (g)

Nematode inocu l ation

3 days beforeDuring 7 dayd after

BABA

Conc.

(mM)

DryFreshDryFresh Dry Fresh

Mean

Dry Mean

Fresh

40 3.2525.321.1513.69 1.92 25.13 2.1021.38

20 2.9022.221.0017.92 1.26 24.06 1.7221.40

10 1.3921.230.5712.88 1.31 14.31 1.0916.14

5 2.2718.340.5313.88 1.24 18.58 1.3516.93

Control1.4017.830.5112.86 1.00 14.48 0.9715.05

Mean2.2420.990.7514.24 1.35 19.31

LSD (P = 0.05) Dry weight: Conc. = 0.2669*, Treat. = 0.2067*, Inter. =

0.4623*. LSD (P = 0.05) Fresh weight: Conc. = 3.20*, Treat. = 2 .4*, Inter. =

5.54*. Each number is a mean of three replicates and two plant each.

*Indicate significant deferens. Nematode inoculums (nematode infested soil

+ peat moss, 1:1) were added when plants were 4 - 5 true leaves . BABA= β

amino butyric acid.

inoculation compared to 3.25 and 4.08 when BABA was

applied during or 7 days after the nematode re spectively.

The lowest gall index, 1.66 was caused by application of

5 mM of BABA 3 days before nematode inoculation

compared with the highest gall index, 5 in nematode in-

fected control.

4.2.2. Effect of Soil Drenching with BABA on

Weight of Roots

Results in Ta bl e 5 showed that soil drenching with 5

S. S. Mutar, F. A. Fattah / Agricultural Sciences 4 (2013) 608-613 611

Table 4 . Effect of soil drenching of BABA on rate of root gall

index caused by Meloidogyne spp on tomato, Solanum lycoper-

sicum L. 30 days after nematode inoculation.

Rate of root gall index

Nematode inocu l ation

BABA Conc.

(mM)

3 days before During 7 days after

Mean

5 1.66 2.33 3 2.33

2.5 2 2.66 3.66 2.77

1 3 3 4.66 3.55

Control 5 5 5 5

Mean 2.91 3.25 4.08

LSD (P = 0.05) Conc. = 0.36*, Treat. = 0.31*, Inter. = 0.62* Each number is

a mean of th ree repli cates and two plan t each. *indicate significant deferens.

Nematode inoculums (nematode infested soil + peat moss, 1:1) were added

when plants were 4 - 5 true leaves. Gall index was according to 1 - 5 level

scale: 1 = no galls on the roots, 2 = galls on 1% - 25% of the root , 3 = galls

on 26% - 50% of the root, 4 = galls on 51% - 75% of the root, and 5 = galls

on 76% - 100% of roots. BABA = β,amino butyric acid.

Ta b l e 5 . Effect of soil drenching of BABA on root weights of

tomato, Solanum lycopersicum L. infected with Meloidogyne

spp 30 days after nematode inoculation.

Root weight (g)

Nematode inocu l ation

3 days before During 7 days after

BABA

Conc.

(mM)

Dry Fresh Dry FreshDry Fresh

Mean

Dry Mean

Fresh

5 0.27 3.25 0.34 3.720.39 9.86 0.355.61

2.5 0.16 2.50 0.39 8.980.51 9.44 0.356.97

1 0.32 3.37 0.49 11.39 0.52 9.43 0.438.06

Control 0.35 3.75 0.36 10.23 0.61 9.77 0.447.91

Mean 0.28 3.21 0.39 8.580.51 9.62

LSD (P = 0.05) Fresh weight: Conc. = 1.41*, Traet. = 1.22*, Inter. = 2.44*.

LSD (P = 0.05) Dry weight: Conc. = 0.0724*, Inter. = 0.1447*. Each number

is a mean of three replicates and two plant each.*Indicate significant

deferens. Nematode inoculums (nematode infested soil + peat moss, 1:1)

were added when plants were 4 - 5 true leaves. BABA = β,amino butyric

acid.

mM BABA significantly (p = 0.05) affected Rfw and

Rdw of Meloidogyne spp infected tomato plants. At this

concentration average Rfw and Rdw were 5.61 and 0.35

g compared with 7.91 and 0.44 g in control plants re-

spectively. However, 2.5 and 1 mM did not significantly

affect root weights. The 40 mM recorded the lowest av-

erage root weights, 5.74 and 0.34 g compared with other

test treatments 30 days after nematode inoculation. Treat-

ment with BABA 3 days before nematode inoculation

produced the lowest average Rfw and Rdw, 3.21 and

0.28 g compared with 8.58, 0.39, 9.62 and 0.51 g at the

time and 7 days after nematode inoculation respectively.

While the lowest root weights were recorded for the 5

and 2.5 mM, 3 days before nematode inoculation, 3.25,

0.27, 2.50, 0.16, the 1 mM concentration and control

recorded the highest root weights, 11.39, 0.49, 10.23,

0.36 g respectively when the treatment was performed at

the timeof nematode inoculation.

4.2.3. Effect of Soil Drenching with BABA on

Weight of Shoots

Soil drenching with the test concentrations of BABA

significantly (p = 0.05) affected shoot weights of Meloi-

dogyne infected tomato plants (Table 6). The average

Sfw and Sdw was grater, 20.43 and 2.34 g in 5 mM

treatment compared with 15.11 and 1.19 g in control.

Grater shoot weights were observed when BABA treat-

ments exceeded nematode inoculation by 3 days , 20.91,

2.65 compared with 13.11, 0.70 and 18.61, 1.52 g when

BABA treatment was at or 7 days after nematode inocu-

lation respectively. The greatest average Sfw and Sdw,

24.32 and 4.22 g were recorded for 5mM application 3

days before nematode inoculation while the lowest Sdw,

0.43 g and Sfw, 11.75 g for the1mM at the time of nema-

tode inoculation and the untreated control plants respec-

tively.

5. DISCUSSION

Results of this work indicated that foliage as well as

soil drench applicatio n of BABA induced systemic resis-

tance against Meloigogyne spp in susceptible tomato

plants. This was manifested by the reduction of galls on

roots and numbers of J2 in roots of BABA treated plants

compared with those in BABA untreated but nematode

Table 6. Effect of soil drenching of BABA on shoot weights of

tomato, Solanum lycopersicum L. infected with Meloidogyne

spp 30 days after nematode inoculation.

Shoot weight (g)

Nematode inocu l ation

3 days beforeDuring 7 days after

BABA

Conc.

(mM)

DryFreshDryFresh Dry Fresh

Mean

Dry Mean

Fresh

5 4.2224.321.0916.47 1.71 20.50 2.3420.43

2.52.7622.920.7511.96 1.64 20.90 1.7118.59

1 1.9020.510.4312.25 1.45 15.39 1.2616.05

Control1.7315.890.5311.75 1.30 17.67 1.1915.11

Mean2.6520.910.7013.11 1.52 18.61

LSD (P = 0.05) Fresh weight: Conc. = 2.50*, Traet. = 2.16*, Inter. = 4.33*.

LSD (P = 0.05) Dry weight: Conc. = 0.319*,Treat. = 0.2763*, Inter. =

0.5252*. Each number is a mean of three replicates and two plant each.

*Indicate significant deferens. Nematode inoculums (nematode infested soil

+ peat moss, 1:1) were added when plants were 4 - 5 true leaves. . BABA =

β,amino butyric acid.

S. S. Mutar, F. A. Fattah / Agricultural Sciences 4 (2013) 608-613

612

infected plants. The growth of the nematode infected

plants by decreasing root weight and increasing shoot

weight of BABA treated tomato plants. These findings

also support previous reports indicating that treatments

with β,amino butyric acid reduced root knot disease

through decreased penetration of J2, gall number on

roots and nematode development [4,11,12]. Root weight

increase in heavy Meloidogyne spp infection was previ-

ously reported and thought to be caused due to biomass

accumulations in infected roots [16]. Meloigogyne spp

infection is known to have negative effects on water and

nutrient elements as well as photosynthesis [17]. It was

reported that M. incognita infection caused biomass ac-

cumulation in roots and this is controlled by the effi-

ciency of the pathogen in capturing the light energy and

directing it in favor of the pathogen or the infected host

[18]. Because of the relative large size of females of

Meloigogyne spp and its ability to produce large number

of eggs, it requires large amount of energy. Beside this

energy requirement, these pathogens caused obvious

distortion in xylem vessels, swellings of root cells and

formation of giant feeding cells which alter root normal

functions. Treatments of nematode infected plants with

the high concentration of BABA produced more shoots

compared with untreated nematode infected plant. This is

mainly due to the fact that Meloidogyne infections em-

bed photosynthesis and chlorophyll synthesis which ne-

gatively influences plant growth [17]. It was clear from

the results of this study that treatments with BABA was

more effective when applied before nematode inocula-

tion, which is probably due to the mechanism of induce

resistance of this chemical inducer. The mechanism of

induce resistance to Meloidogyne in tomato by BABA is

not fully understood. It was believed that treatments with

this inducer render roots less attractive to J2 through al-

tered plant nutrient assimilation or render plant cell walls

harder to penetrate by J2 or that giant cells were smaller

or not able to provide enough nutrients for the develop-

ing nematodes [11]. Treatments with BABA were re-

ported to increase levels of salicylic acid (SA) and patho-

genesis related proteins (PRP) [19], and enzymes like

catalase (CAT), polyphenoloxidase (PPO) and guaiacol

peroxidase (GPOX) [4,7] and phenol compounds [20].

BABA was also reported to induce the accumulations of

PPO, GPOX, H2O2, CAT and phenols in M.javanica in-

fected cucumber roots [21].

Results of this research work clearly indicate and in-

dorse previous reports on the effectiveness of β,amino-

butyric acid to the induced systemic resistance as envi-

ronmentally sound alternative for the control of Meloi-

dogyne sp p on t omato.

6. ACKNOWLEDGEMENTS

The authors would like to thank Mr. Dhulfiqar Layth (Graduate stu-

dent, Dept., of Plant Protection, College of Agriculture,Univ. of Bagh-

dad, Baghdad, Iraq) for his valuable assistance.

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