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<strong>SCREENING</strong> <strong>OF</strong> <strong><strong>TO</strong>MA<strong>TO</strong></strong> <strong>ACCESSIONS</strong> <strong>FOR</strong> <strong>RESISTANCE</strong> <strong>TO</strong> FUSARIUM WILT<br />

CAUSED BY Fusarium oxysporumj.sp. Iycopersici IN SOUTHWEST Nigeria<br />

A PROJECT SUBMITTED <strong>TO</strong> THE, DEPARTMENT <strong>OF</strong> CROP PROTECTION,<br />

COLLEGE <strong>OF</strong> PLANT SCIENCE AND CROP PRODUCTION UNIVERSITY <strong>OF</strong> .<br />

AGRICULTURE, ABEOKUTA, OGUN STATE, NIGERIA.<br />

IN PARTIAL FULFILMENT <strong>OF</strong> THE REQUIREMENTS <strong>FOR</strong> THE DEGREE <strong>OF</strong><br />

BACHELOR <strong>OF</strong> AGRICULTURE (B.AGIUC), <strong>OF</strong> THE UNIVERSITY <strong>OF</strong><br />

AGRICULTURE, OGUNSTATE, NIGERIA.


DEDICATION<br />

This project is dedicated to God Almighty, who has been my strength and helper throughout<br />

my<br />

undergraduate<br />

days in UNAAB.


My gratitude goes to our Lord Jesus Christ, the greatest provider, sustenance and giver of<br />

peace.<br />

My parents; Mr and Mrs. Oduyeru, I really appreciate you for your support, morally and<br />

financially,<br />

may you live long to eat the fiVit of your labour in Jesus name, Amen. My siblings<br />

Miss Temitope, Damilola, Busola, BiodUIl: and Segun, My Daddy (Mr Wale LawaI) for your<br />

fatherly support and Advice and my Big Uncles; Mr Deji Lawal, Mr Ranti Oduyeru and Mr<br />

Gboyega Okubanjo thanks for your understanding<br />

at all time.Also, to all_my immediate relatives,<br />

thanks for your support. We shall all live to see more of His goodness.<br />

To my supervisor: Dr. A.R Popoola, .thank you for your fatherly advice and for all the<br />

knowledge<br />

you imparted in me. I am really grateful. And not forgetting Mr. Ganiyu for your<br />

Advice and moral support throughout this project<br />

To my Head of Department;<br />

Dr Pitafi And to my lecturers, Dr. Atugun, Mr. Odeyemi, Dr.<br />

Ayo-John and Dr. Afolabi and to others that their names are not mentioned here, thank you sirs.<br />

To my project mates, in person of Lekan Davies, Kuponiyi Razak and Bimbola Adefolalu., God<br />

bless you all for your kind gesture all thr01,lgh. And not forgetting my friend and a brother from<br />

another mother in person of Adami Ayokunle, thanks a million. May our labour never be in vain.


The experiment<br />

was conducted to investigate the resistant level of tomato accessions to Fusarium<br />

wilt caused by Fusarium oxysporum f sp.Lycopersici. The experiment was a randomized<br />

complete block design (RCBD) consisting of 3 treatments with 3 replicates. Ten accessions were<br />

inoculated with samples of fungal strain 1 and strain 2 using Drenching method into the soil.<br />

Plant performance<br />

was based on plant height, number of leaves number of branches and number<br />

of tomato fruits. Disease assessment was based on disease severity, incidence and number of<br />

days for 50% wilt. The result obtained from the experiment shows that different variety has<br />

different resistant level to the disease which was evident in the plant height and number of fruits<br />

produced.NG/MRIMA Y/09/006 is resistant to strain 1 and susceptible to strain 2 of the Fusarium<br />

been inoculated into it and was able to produce the high number of fruits.<br />

This study could therefore suggest N GIM RIMA Y109/006 which has minimum mean severity<br />

(36.03%) and could produce reasonable number of fruit under the Fusarium wilt infection.


1.0 Introduction<br />

1.1 Tomato<br />

1.2 Botanical description<br />

1.3 Disease resistance<br />

1.4 Abiotic stress tolerance<br />

1.5 Objective<br />

1.6 Justification


1: Disease Severity scale<br />

2: Disease assessment oftomato accessions under infection and control<br />

3: Growth performance of tomato accessions.<br />

4: Yield parameter of Tomato accessions


Tomato (Lycopersicon escuientum Mill) is an important source of vitamin C,<br />

calories, phosphorus and calcium (Davis and Hobson, 1981; Langer and Hill, 1991).Worldwide,<br />

tomatoes are an important part of a diverse and balanced diet (Willcox et ai., 2003). It is estimated<br />

that over 62 800 accessions of the cultivated and wild species of tomato (mostly L. escuientum<br />

accessions) are maintained in genebanks around the world(Rossi et ai., 1998), Among the nine<br />

Lycopersicon species, only" L. escuientum has become a domesticated crop though L.<br />

pimpinellifolium (with fruit diameter -1 cm) is also casually planted for consumption(Rick,<br />

1978).The cultivated tomato has limited variability, largely because of several population<br />

bottlenecks in the forms of founder events and natural and artificial selections that occurred during<br />

domestication and evolution of modem cultivars(Rick, 1976),During the past 70 years wild species<br />

of tomato have been utilized in breeding programs to improve the cultivated tomato (Rick,.<br />

1979;.Rick, 1982), According tos (Rick, 1976), domestication of L. escuientum<br />

was accompanied<br />

by a transition from exerted to inserted stigmas, and consequently changing from facultative<br />

outcrossing to enforced inbreeding, It is not known exactly -when domestication of tomatoes<br />

occurred, however, by the time the Spanish conquered Mexico in 1523, they were already<br />

domesticated (Rick, 1978).


1.2 Botanical description<br />

Tomato belongs to the nightshade family Solanaceae, which is in division<br />

Magnoliophyta, class Magnoliopsida, subclass Asteridae, order Solanales, and suborder Solanineae<br />

(Knapp et al., 2004; Nee et al., 1991.). The extremely diverse and large Solanaceae family is<br />

believed to consist of 96 genera and over 2800 species in three subfamilies, Solanoideae (in which<br />

Lycopersicon belongs), Cestroideae, and Solanineae. Solanaceae is one of the most economically<br />

important families of angiosperms and contains many of the commonly ~ultivated plants, including<br />

potato, tomato, pepper, eggplant, petunia, and tobacco. This family is the most variable of all crop<br />

species in terms of agricultural utility, the 3rd most economically important crop family, exceeded<br />

only by the grasses and legumes, and the most valuable in terms of vegetable crops (Vander<br />

Hoeven, et al., 2002). Among all plant families, members of the Solanaceae are extremely diverse in<br />

terms of growth habit (from trees to small annual herbs), habitat (from deserts to the wettest tropical<br />

rain forest), and morphology (Knapp et al., 2004). Many Solanaceous species have played important<br />

roles as model plants, including tomato, potato, pepper, tobacco, and petunia. The tomato genus<br />

Lycopersicon is one of the smallest genera in Solanaceae, though the centerpiece in the family for<br />

genetic and molecular research. It is the closest to the genus Solanum (nightshade), an association<br />

which originally led people to believe tomato was poisonous (Rick, 1978). The cultivated tomato<br />

was originally named Solanum Iycopersicum by Linnaeus.(Linnaeus, 1953).<br />

Tomato is susceptible to over 200 diseases caused by pathogenic fungi, bacteria,<br />

viruses, or nematodes (Lukyanenko, 1991). Without question, the greatest contribution of modem<br />

plant breeding to tomato improvement has been through development of cultivars with improved<br />

disease resistance. Resistance has been identified, and in many cases characterized, for more than 30


of the major tomato diseases. Most commercial cultivars possess up to 6 (in true breeding lines) or<br />

10 (in hybrids) disease-resistance attributes. These mainly include diseases for which major<br />

resistance genes have been identified, including fusarium wilt, verticillium wilt, root-knot nematode,<br />

alternaria stem canker, gray leaf spot, and some bacterial and viral diseases. However, horizontal<br />

(a.k.a. field or polygenic) resistance has also been reported for several tomato diseases, where major<br />

genes for resistance to a particular pathogen or race are not found, such as early blight, powdery<br />

mildew, bacterial canker, and bacterial wilt. Except in a few cases, tomato wild species have been<br />

utilized as the source of resistance for all tomato diseases. Resistance resources have been identified<br />

in most related wild species of tomato, in particular 1. pimpinellifolium, 1. peruvianum, and 1.<br />

hirsutum. For some tomato diseases, such as late blight (caused by oomycete Phytophthora<br />

infestans) and powdery mildew (caused by fungus Oidium lycopersicum), both vertical and<br />

horizontal resistances have been identified.<br />

Although the cultivated tomato is widely adapted to different climates, its growth and<br />

development is rather sensitive to different environmental stresses, including salinity, drought,<br />

excessive moisture, extreme temperatures, mineral toxicity and deficiency, and environmental<br />

pollution. There is limited genetic variation for abiotic stress tolerance within the cultivated species<br />

and most commercial cultivars are considered moderately to highly sensitive to different stresses.<br />

Fortunately, sources of genetic tolerance (or resistance) to different abiotic stresses are found within<br />

the related wild species, including 1. chilense, 1. peruvianum, 1. pennellii, 1. pimpinellifolium, 1.<br />

hirsutum,1. cheesmanii, 1. chmielewskii, and 1. parviflorum (Foolad, 2005). In addition, there are a<br />

few species within Solanum that exhibit tolerance to environmental stresses and which may be<br />

utilized in tomato breeding for stress tolerance. Several tomato wild species have been utilized for


genetic and physiological characterization of abiotic stress tolerance and for breeding purposes<br />

(Scott, 1993), In addition, stress tolerance is a developmentally regulated, stage-specific<br />

phenomenon; tolerance at one stage of plant development is often not correlated with tolerance at<br />

other developmental stages (Asins et al., 1993; Foolad, 1999).<br />

1.5 Objective:<br />

To determine the resistance levels of different tomato varieties to Fusarium wilt<br />

disease caused by Fusarium oxysporum f<br />

sp. Iycopersici.<br />

1.6 Justification:<br />

Tomato varieties are susceptible to several diseases caused by pathogenic fungi,<br />

bacteria or viruses. Different c?ntributions of modem plant breeding to tomato improvement have<br />

been through development of cultivars with improved disease resistance. Hence the need to<br />

determine the resistance level of different varieties of tomato to Fusarium caused by Fusarium<br />

oxysporumfsp<br />

lycopersici wilt disease in South West Nigeria.


· .<br />

plant roots, but seem not to invade the vascular system (Olivain and Alabouvette 1997, 1999). The


However, there are indications for a certain F: oxysporum-strain<br />

specific effect of compounds<br />

found in tomato root exudates. In a recent study it was reported that the germination rate of several<br />

F. oxysporum strains is affected differently by tomato root exudates (Steinkellner et al., 2005).<br />

Moreover, studying the spore germination of a Fori strain and a biocontrol strain of F. oxysporum in<br />

tomato root exudates and two of its major known compounds, glucose and citric acid, (Bolwerk et<br />

al., 2005) found a higher stimulation for the biocontrol strain than for the pathogenic strain. In<br />

several studies it has been shown that root colonisation by symbiotic arbuscular-mycorrhizal fungi<br />

alters the root exudation pattern and thus the effect of the exudates on soil-inhabiting microbes<br />

(Lioussanne et al., 2003; Pinior et al., 1999; Sood, 2003). (Pinior et al., 1999) reported that root<br />

exudates from mycorrhizal plants affect the hyphal growth of arbuscular mycorrhizal fungi<br />

differently, compared to root exudates from non-mycorrhizal plants. Recently, (SchefIknecht et al.,<br />

2006, 2007) reported that root colonisation by an arbuscular-mycorrhizal fungus also affects the<br />

microconidial germination of the tomato pathogen F. oxysporum.<br />

2.2 Biology of Fusarium<br />

In solid media culture, such as potato dextrose agar (PDA), the different special forms<br />

of F. oxysporum<br />

can have varying appearances. In general, the aerial mycelium first appears white,<br />

and then may change to a variety of colors - ranging from violet to dark purple - according to the<br />

strain (or special form) of F. oxysporum.<br />

If sporodochia are abundant, the culture may appear cream<br />

or orange in color (Smith et al., 1988).<br />

F. oxysporum produces three types of asexual spores: microconidia, macroconidia, and<br />

chlarnydospores (Agrios, 1988).Microconidia are one or two celled, and are the type of spore most<br />

abundantly and frequently produced by the fungus under all conditions. It is also the type of spore


most frequently produced within the vessels of infected plants. Macroconidia<br />

gradually pointed and curved toward the ends. These spores are commonly<br />

are three to five celled,<br />

found on the surface of<br />

plants killed by this pathogen a


Due to the growth of the fungus within the plant's vascular tissue, the plant's water supply is greatly<br />

affected. This lack of water induces the leaves' stomata to close, the leaves wilt, and the plant<br />

eventually dies. It is at this point that the fungus invades the plant's parenchymatous<br />

tissue, until it<br />

finally reaches the surface of the dead tissue, where it sporulates abundantly (Agrios, 1988). The<br />

resulting spores can then be used as new inoculums for further spread of the fungus<br />

Due to prolonged survival in soil as a saprophyte and as resistant structures, F<br />

oxysporum is difficult to control (Mujeebur and Shahana, 2002; Borrero et al., 2004). Among control<br />

strategies available for Fusarium wilt management, soil fumigation and use of resistant cultivars are<br />

the most used. Biological control of plant pathogens using antagonistic<br />

fungal and bacterial strains is<br />

gaining increasing importance. Soil has untapped potential and contains several potential biocontrol<br />

agents (BCAs) such as Pseudomonas spp., Trichoderma spp. and non-pathogenic Fusarium spp. that<br />

have shown high antagonistic activity against several soil-borne pathogens (Fuchs et al., 1999;<br />

Ramamoorthy et aI., 2001). So far, several bacterial and fungal antagonistic strains have been shown<br />

to reduce the damage caused by different pathogens, including F oxysporum f. sp. Iycopersici<br />

(Postma and Rattink, 1992; Alabouvette, 200 I; Moretti et al., 2008).<br />

The best recommended<br />

way to control the disease is selecting resistant varieties of tomato (Silva and<br />

Bettiol, 2005). Recently there have been many reports about using bioactive compounds from<br />

biological control agents which were extracted from different fungi that inhibit many plant<br />

pathogenic fungi, including Fusarium wilt of tomato (Soytong, 1992). These bioactive compounds<br />

are Tricotoxin A50 extracted from Trichoderma harzianum PCO1; and Chaetoglobosin C extracted


from Chaetomium globusum that have been reported to elicit resistance or immunity in plants by<br />

inducing oxidative burst in plant cells (Nuchdonrong, et al., 2004; Soytong, 2002).<br />

So far, several bacterial and fungal antagonistic strains have been shown to reduce the damage<br />

caused by different pathogens, including F. oxy:sporum f. sp. Iycopersici (Postma and Rattink, 1992;<br />

Alabouvette, 2001; Moretti et al., 2008). Some studies have shown that the combined effect of<br />

antagonism and plant growth promotion by BeAs suppresses Fusarium wilt disease in several crops<br />

(De Boer et aI., 2003; Fahri and Murat, 2007).


The following ten accessions of tomato were used for the study:<br />

.


Dextrose Agar (PDA) and incubated at room temperature with the morphological<br />

characteristic of Fusarium been studied under compound microscope. F. oxysporum after<br />

incubation was purified by repeating sub-culture into fresh sterile media until pure culture<br />

were examined on the microscope. Propa!?ules was prepared in 100ml distilled water. The<br />

resultant suspension, with a concentration of 10 6 _10 7 propagules per ml was used to inoculate<br />

tomato plants 2wks after transplanting.<br />

3.5 Screenhouse experiment:<br />

The experiment was in completely randomized design with 3 replicates<br />

comprising of 90 space pots of accession's population maintaining intra and Inter spacing of<br />

0.5m and 1m within the pots. Platform comprising of bamboo were made,occupied by 10m<br />

by 5m land area space in which the pots was placed to avoid soil borne pathogen into the pot<br />

experimental<br />

pots.<br />

Cultural practices: wetting and weeding were regularly observed.<br />

Trans plan ting:<br />

Tomato seedlings of 4 wks old were transplanted into pots containing sterilized<br />

Tomato accessions = 10<br />

Fusarium strains = 2 most virulent + 1 control = 3


All data were subjected to analysis of variance (ANOYA) and mean were separated DMRT<br />

(Duncan Multiple Range Test)


% range scale Value<br />

1-20.0 1 Resistant (R)<br />

20.01-40.00 2 Mean resistant (MR)<br />

40.01-60.00 3 Mean susceptible (MS)<br />

60.01-80.00 4 Susceptible (S)<br />

~0.01-100.00 5 Highly susceptible (HS)<br />

Kavak and Boydak, 2006


The'result of the effect of Fusarium wilt on agronomic and yield parameters of the<br />

tomato accessions are presented in Table 2, 3, and 4. The number of branches, plant height and<br />

number of leaves per plant recorded in this study were as a result of their genetic resistance. The<br />

resistant accessions maintained their leaf numb~rs despite the Fusarium,wilt.<br />

Based on the number of leaves and plant height on Table (3), NG/MRIMA Y/09/005<br />

has 53.68cm for Strain 1 and 51.78cm for strain 2 while NG/MRIMAY/09l006<br />

has 54.88cm for<br />

strain 1 and 65.89cm for strain 2 and are more resistant to both strains of Fusarium inoculated and<br />

the accession that recorded the highest number of the leaves/plant is NG/MRIMA Y/09/006.<br />

Yield parameters such as number of fruits harvested were considered important<br />

indicators for the Fusarium wilt Table (4), in this study, the tomato fruits were harvested when they<br />

were firm and their colour fully developed. The number of fruit Iplant of the control is higher than<br />

the infected except in some cases which could be as a result of environmental factors, thus the<br />

Fusarium affects the number of fruitiplaI1t.NGIMRlMAY/09/006<br />

produces the highest number of<br />

fruit under infection, with strain 1 with having 4 and strain 2 with 2.67<br />

The incidence was compared among the varieties during the study on Table (2), both<br />

partially and incompletely wilted plant were encountered on the field, the result shows that the<br />

incidence of fusarium wilt on tomato is high in NGIMRIMA Y/09J005, having 3.50 to strain 1 and<br />

3.00 to strain 2 while NGIMRIMA Y/09/006 shows minimum resistance to the infection, having 2.33


to strain 1. A common feature on the fusarium wilt -on plant were collected from thO<br />

presence of varying degree of chlorosis.


ROMAROUND 2.67 ab 2.00 a 3.33 ab 45.03 ab 30.85 a 51.26 ab 48.33 a 45.00 a 46.00 a<br />

BESKE 2.67 ab 3.00 a 3.33 ab 45.00 a 44.83 a 52.13 ab 47.67 a 46.67 a 43.00"<br />

NG/M R/MAY/09/005 3.50 b 3.00" 2.67"b 57.10 b 46.00" 44.10 a 40.50" 47.33 a 52.00"<br />

NG/MR/MAY/09/006 2.33 ab 3.00" 4.00 b 36.76" 41.30 b 63.03 b 53.33 a 52.33 a 42.33 a<br />

NG/OE/MAR/09/019 2.67"b 3.00 a 3.33 ab 41.1 a 53.30 ab 49.53" 46.67" 44.67" 46.33 a<br />

PUREWATER 3.00"b 2.00 a 3.67 ab 53.65 a 33.75 b 64.06" 44.50" 56.00" 38.00 a<br />

SANTANA 2.00 a 1.50" 2.33 a 31.67" 26.00" 41.70"b 57.00" 60.50 a 53.67"<br />

NG/AO/MAY/09/011 2.00" 3.00 a 2.33 a 33.13 a 53.33"b 37.03" 56.00 a 44.33" 52.33 a<br />

NG/AA/MAY/09/030 3.50 b 2.33 a 3.00"b 53.35 ab 38.13" 48.60 ab 43.00 a 50.67" 46.33 a<br />

NG/TB/ AUG/09/006 2.00 a 3.00 ab 3.00 ab 48.80 a 48.80 a 49.83" 54.33" 45.67" 43.67"<br />

Means follow by a common letter in each column are not significantly difference by DMRT at P =<br />

0.05


TABLE 3: Growth performance of tomato accessions<br />

Accessions Plant height Number of Leaves number of branches<br />

51 52 CONTROL 51 52 CONTROL 51 52 CONTROL<br />

ROMAROUND 47.39" 31.96"b SS.40 b 8.22" 8.66 b 8.77" O.SO"b 0.15" 0.25"<br />

BESKE 35.55" 45.23" 60.S0~ 6.33" 7.89 b 8.44" 0.67" 0.25" 0.67"<br />

NG/MR/MAY/09/005 53.68" 51.78 bc S4.70 b 7.94 b 7.00 b 9.11" 0.33" 0.33" 1.00 b<br />

NG/MR/MAY/09/006 54.88" 65.89" Sl.30"b 7.56" 10.89" 8.67 b 0.33" 1.67"b 0.7S"b<br />

NG/OE/MAR/09/019 37.48" SS.OSb Sl.78"~ 6.00" 11.00" 7.SS b 1.00" 0.67"b 0.67"<br />

PUREWATER 38.76" 35.13"b 46.38"b 7.00 b 6.5S"b 7.5S b 0.33" 0.50" 1.00"b<br />

SANTANA 37.73" 25.23" 36.76" 7.44 b 5.67" 8.22" 0.25" 0.45" 1.00"b<br />

NG/AO/MAY/09/011 47.21" 44.07"b 38.93" 7.4S" 6.67"b 5.77" 0.33" 0.39" 0.33"<br />

NG/AA/MAY/09/030 41.15" 44.53"b 51.32"b 6.78 b 6.67"b 8.00" 0.45" 0.36" 0.33"<br />

NG/TB/ AUG/09/006 40.72" 44.76"b 44.6S"b 6.33 b 6.77 b 7.44" 0.35" 0.19" 0.33"<br />

Means follow by a common letter in each column are not significantly difference by DMRT at P =<br />

0.05


Table 4: Yield parameter of tomato accessions<br />

<strong>ACCESSIONS</strong> .<br />

Number of fruits<br />

$1 52 CONTROL<br />

ROMAROUND 2.67 3 0.50 3 2.67 3<br />

BESKE 1.00 3 2.00 3 3.33 3<br />

NG/MR/MAY/09/00S 1.33 3 2.00 3 3.33 3<br />

NG/MR/MAY /09/006 4.00 3 2.67 3 2.33 3<br />

NG/OE/MAR/09/019 0.67 3 2.00 3 1.67 3<br />

PURE WATER 1.33 3 0.67 3 2.33 3<br />

SANTANA 1.00 3 0.00 3 1.67 3<br />

NG/ AO/MAY /09/011 1.67 3 2.67 3 0.67 3<br />

NG/ AA/MAY /09/030 0.00 3 1.00 3 2.33 3<br />

NG/TB/ AUG/09/006 0.67 3 2.00 3 1.33 3<br />

Means follow by a common letter in each column are not significantly difference by DMRT at P =<br />

0.05


The tomato accessions were found to vary in their resistance to strain 1 and strain 2 of<br />

Fusarium oxysporumf<br />

sp. Lycopersici .This study shows that some diversity exists in the tomato<br />

accessions.<br />

Severity test for strain 1 shows that 7 varieties showed mean resistance while<br />

NGIANMAY/09/030, Pure wate~ and NG/MRIMAY/09/005 were susceptible. Maximum rating was<br />

3.5 on NG/MRIMAY/09/005 and NGIANMAY/09/030 when challenged with stain 1 on the field.<br />

Under field condition, many of the varieties that were grown were severely affected. Almost all<br />

varieties were susceptible to strain 2 of the inoculated Fusarium<br />

It is well documented<br />

that the occurrence of disease in plant was found to depend on at<br />

least two components: pathogen and its host plant; and for disease development requires favorable<br />

environmental<br />

condition. This study showed the different resistant level of the ten accessions to<br />

Fusarium wilt disease which is a soil borne disease.<br />

NG/MRIMA Y/09/006 exhibited minimum resistance in terms of disease severity,<br />

number of leaves and fruit yield, with its yield comparably<br />

high, which makes the variety as the<br />

best performing accessions used. It was observed from the data taken that all the tomato<br />

accessions inoculated with the fungi, fusarium oxysporum fsp .lycopersici shows the symptoms<br />

of Fusarium wilt, therefore the best way to control the disease is by planting under irrigation<br />

during the dry season


Based on this study, NG/MR/MA Y/09/006 is recommended in Fusarium wilt endemic<br />

field and also recommended is NG/AO/MAY/09/011 which is recommended based on high yield<br />

and the level of resistant to Fusarium wilt exercised by these varieties.


·<br />

different disease suppressive mechanism. Phytopathology 93:626-632


Fahri, Y., Murat D. (2007) Control of Fusarium wilt of tomato by combination<br />

of Fluorescent<br />

Pseudomonas, nonpathogen Fusarium and Trichoderma hazianum T22 in greenhouse<br />

condition. Plant Pathology Journal 6: 159-163.<br />

Foolad M.R. 1999, "Comparison of salt tolerance during seed germination and vegetative growth in<br />

tomato by QTL Mapping, "Gssenome. vol.42 no. 4, pp.727-734<br />

Foolad M.R. 2005, Breeding for abiotic stress tolerance in tomato," in Abiotic Stresses: Plant<br />

Resistance Through Breeding and Molecular Approaches, Ashraf M, and Harris P.J.C, Eds.,<br />

pp. 613-684, The Haworth Press, New Yor1


Olivain C. And Alabouvette C. (1999) Processes of tomato root colonization by a pathogenic strain of<br />

Fusarium oxysporum f.sp. lycopersici in comparison with a non-pathogenic strain. New<br />

phytologist, 141,497-510<br />

Olivain, C., and Alabouvette C. (1997) Colonizations of tomato root by a non-pathogenic strain of<br />

Fusarium. New Phytologist, 137,481-494.<br />

Olivain, C.,Humbert C.,Nahalkova J.,Fatehi J.,L'haridon F., and Alabouvette C. (2006).Colonization of<br />

tomato root by pathogenic and nonpathopgenic Fusarium oxysporum strains inoculated together<br />

and separately into the soil. ,Applied and Environmental Microbiology, 72, 1523-1531<br />

Pinior, A., Wyss D., Piche Y., and Vierheilig H. (1999) Plants colonized by AM fungi regulate further<br />

root colonization bt AM fungi through altered root exudation. Canadian Journal of Botany,<br />

77,891-897<br />

Postma J., Rattink H., 1992. Biological control of Fusarium wilt of carnation with a non pathogenic<br />

isolate of Fusarium oxysporum.<br />

Canadian Journal oj Botany 70: 1199-1205; Potato and<br />

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