A new source of resistance to bacterial wilt of eggplants obtained from a cross : Solanum aethiopicum L. X Solanum melongena L

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A new source of resistance to bacterial wilt of eggplants

obtained from a cross : Solanum aethiopicum L. X

Solanum melongena L

G. Ano, Philippe Prior, C.M. Messiaen

To cite this version:

G. Ano, Philippe Prior, C.M. Messiaen. A new source of resistance to bacterial wilt of eggplants

obtained from a cross : Solanum aethiopicum L. X Solanum melongena L. Agronomie, EDP Sciences,

1991, 11 (7), pp.555-560. �hal-02716116�

Plant

_pathology

A

new

source

of resistance

to

bacterial wilt

of

_eggplants

obtained from

a

cross:

Solanum

_aethiopicum

L

x

Solanum

melongena

L

G Ano

Y Hebert

P Prior

CM

Messiaen

1 _{INRA CRAAG, Station d’Amélioration des}

Plantes, BP 1232, 97184 Pointe-à-Pitre Cedex;

2

INRA, station d’Amélioration des Plantes Fourragères, 86600 Lusignan;

3_{INRA CRAAG, Station de}

Pathologie Végétale

et de

_{Malherbologie,}

BP 1232, 97184 Pointe-à-Pitre _{Cedex, Guadeloupe;}

4

INRA, Station de _Biologie et de

_{Pathologie Végétale,}

Place Viala, 34060

_Montpellier

Cedex, France

(Received 3

_May

1990;

accepted

29 _{April 1991)}

Summary —

As the

_{eggplant hybrid}

_F

₁

Kalenda no

_longer

showed

_enough

resistance to Pseudomonas

solanacea-rum EF Smith in

_Guadeloupe

and

_{Martinique (French}

West

_Indies),

a new

_{breeding program}

was undertaken

_using

Solanum

_aethiopicum

as source of resistance.

_{Despite sterility problems}

encountered

_during

the first

_generations,

se-lection was conducted under an artificial inoculation test

_according

to a recurrent selection scheme

_including

back-crosses

_by

Solanum

_melongena.

From the second backcross it was

_possible

to obtain families with a

_high

level of

re-sistance to bacterial

_wilt,

as well as a wide variation in the

_shape

and colour of the fruits.

Solanum

_melongena

/ Solanum

_aethiopicum

/ Pseudomonas solanacearum / resistance / recurrent selection

Résumé — Une nouvelle source de résistance au flétrissement bactérien obtenue à

_partir

d’un croisement

entre Solanum

_aethiopicum

L et Solanum

_melongena

L. Un nouveau _programme de sélection

_{d’aubergines}

utili-sant Solanum

_aethiopicum

L comme source de résistance à Pseudomonas solanacearum EF Smith a été lancé en

Guadeloupe

et en

_{Martinique; l’hybride F}

₁

Kalenda ne manifestant

_plus

une résistance suffisante. La sélection a pu être réalisée

_{malgré d’importants problèmes}

de stérilité rencontrés

_pendant

les

_{premières générations.}

Elle fut

con-duite

_{généralement}

sous inoculation artificielle selon un modèle de sélection récurrente avec des

_phases

de

back-cross _par Solanum

_melongena

L.

_Après

le second

_backcross,

il fut

_possible

de sélectionner des familles manifestant

un haut

_degré

de résistance à la bactérie et

_possédant

une très

_large

variabilité pour la forme et la coloration du fruit.

Solanum

_melongena

/ Solanum

_aethiopicum

/ Pseudomonas solanacearum / résistance / sélection récurrente

INTRODUCTION

Bacterial wilt caused

_by

Pseudomonas

solanac-earum EF Smith

_(Kelman,

₁₉₅₃₎

leads

to

important

losses in solanaceous crops in

Martinique

and

_{Guadeloupe (French}

West

In-dies,

Carribean

_{area) (Messiaen, 1975).}

As

in

other

_tropical

and

_{subtropical regions}

of the

world

_{(Buddenhagen, 1985),}

this

disease

is

a

limiting

factor to

the

_development

of

_eggplant

cultivation.

A

_breeding

_{program conducted}

_by

_Daly

(1972,

1973) produced

the line

Madinina

with a

good

level of resistance to the bacterium.

Then,

fruit

anthracnosis,

a new

disease caused

_by

Colletotrichum

_{gloeosporioïdes}

P

_(Fournet,

1973)

led

Kaan

₍₁₉₇₃₎

to

breed

a new

_variety,

Aranguez,

which was resistant to anthracnosis.

The

well

_shaped

and coloured

_variety

was

crossed with Madinina

in

order to

_produce

the

F

1

hybrid,

Kalenda

_{(Daly, 1986).}

Kalenda

is

highly

resistant

to

anthracnosis and shows

good

tolerance

to bacterial wilt.

*

In

_1982,

_damage

caused

_by

P solanacearum

led

to

extensive losses

in

_{eggplant plantations}

on

both

islands

_{(Messiaen, 1983).}

The

_F

₁

_hybrid

Kalenda, still

cultivated

_today,

did not

show

suffi-cient resistance

to

bacterial

wilt,

_especially

in

conditions such

as

hot

season

_planting

(July-August) (Bereau

and

_Messiaen,

₁₉₇₅₎

or

plant-ing

in

_poorly

drained

fields.

Hebert

₍₁₉₈₅₎

showed

that

a

high

level of

re-sistance to

bacterial wilt is available in the

Sola-num

_aethiopicum

_{Aculeatum group. This}

_species

is

often

described under

the name

S

integrifoli-um

_(Lester

and

_Niakan,

_{1986). Beyries (1979)}

used it as rootstock for

_eggplant

and tomato in

naturally

infected fields.

Until now, S

_{aethiopicum species}

as a source

of

bacterial wilt

resistance has

_only

been used

in the cross

with S

melongena

because of

_infertility

problems

in order to obtain

_F

₁

_hybrids

used as

rootstock

_{(Sakaï, 1984).}

This paper

reports

on the first

_phase

of

the

breeding

program

conducted

at INRA

_(French

West

_Indies)

in order to transfer

the

S

aethiopi-cum Aculeatum

_group’s

resistance to

P

solanac-earum

into commercial

_eggplant

varieties.

MATERIALS

AND

METHODS

Inoculation

_techniques

Since our

_objective

was to find a better resistance lev-el than that of

_Kalenda,

this

_variety

was chosen as the

susceptible

control. Two

_techniques

were used: infect-ed

field,

in order to test the

_progenies

of the crosses with the

_{susceptible parent Aranguez;}

and artificial

in-fection,

to

_identify

resistant families in the

_following

generation.

Infected

field

Trial

_plots

were used which had received an artificial inoculation 1 or 2 _years earlier. Under these

condi-tions,

the inoculum is considered to be

homogeneous-ly

distributed over the

_plot

and

_selecting

_{pressure is}

in-termediate.

Artificial infection

So that all the

_plants

could be

_exposed

to the same

strong

infectious

_conditions,

_screening

for resistant

plants

was

_{completed according}

to the method de-scribed

by

Prior et al

_(1989a).

This method is based

on artificial inoculation with a mixture of 3 strains

(GA1,

GA3,

GA4)

isolated from

_eggplant.

These strains were collected from different locations in

Guadeloupe

and have been

_deposited

in the Collec-tion NaCollec-tionale de Bactéries

_{Phytopathogènes}

(CNBP,

INRA Station de

_{Pathologie Végétale,}

49000,

Angers,

France).

All these strains

_belong

to Race 1

(Budden-hagen et al, 1962)

and Biovar III

_{(Hayward, 1964),}

but

they

differ in _{host range}

_(Prior

and

Steva,

1989)

and in

aggressivness (Prior

et

al,

1990).

Virulent fluidal

_type

colonies of the bacterium were _grown on tetrazolium chloride

_(TZC)

medium

_{(Kelman, 1954).}

After 48 h in-cubation at

30 °C, single

colonies were cultivated for

18 h in

_liquid

TZC-free medium at

_{30 °C,}

then harvest-ed and rinsharvest-ed with sterile distillharvest-ed water

_by

centrifuga-tion

_{(4 000}

_g, 20

_min).

Final concentration of the mixed

suspension

was

_{optically adjusted}

to 2 x 107 CFU/ml. At

_planting

time

_{(60 days}

after

_sowing)

2 ml of

inocu-lum were

_placed

close to the roots of each

plant.

All the tested families were

_planted

in a randomized

complete-block design

with 4

_{replicates (15 plants per}

replicate).

Considering

that in the case of

_eggplant

a wilted

_plant

dies

_{rapidly, percentages}

of dead or wilted

plants (WP)

were observed 70

_days

after inoculation.

Breeding

for

resistance

to P

solanacearum

(fig 1)

In 1982, several crosses were made between different S

_aethiopicum

entries from the INRA collection

(Sta-tion d’Améliora(Sta-tion des Plantes

Maraîchères,

84140

Montfavet, France)

used as female

_parents

and

tropi-cal varieties of S

_{melongena. Embryo}

culture with V3 medium

_(Schoch

and

Sibi,

1978)

was used in order to

obtain more viable

_F

₁

_plants

from these crosses.

Inter-specific hybrids

were crossed with

_Aranguez.

A _very

low

_fertility

level was observed

during

this first

_phase

of the _{program, and only} the

progenies

of the cross,

named

_{BPA, —}

_{(S aethiopicum}

ref Bot 2 x S

melonge-na cv

_P18)

x

_{Aranguez -}

were obtained in 1983.

P18,

a line of Turkish

_origin,

has a

_good

resistance to fruit anthracnosis and bacterial wilt

_{(Vincourt, unpublished}

observations)

but its fruit color is not suitable because of the sun’s influence on the

_{purple shade. The}

proge-nies of the BPA cross were allowed to

_{open-pollinate}

for 2

_generations.

At the end of

_1984,

this material had recovered a better

_though

not total

fertility.

Non-viable combinations had been eliminated.

Later on, recurrent selection with backcrosses was

performed.

Selection and

_{intercrossing}

were followed

by

a backcross

_phase

with

_Aranguez

in order to im-prove the quality of the fruit

_{(colour, shape, size).}

After the 2

generations of open

pollination,

equiva-lent to a

_{polycross phase,}

fruits from 41

_plants

result-ing

from the BPA cross were harvested. In November

1984,

the

_offsprings

(named

FAM

_families)

of these 41

plants

were screened

_{by artificially}

_inoculating

_plants

with P solanacearum.

_Only

36 _{families gave} a suffi-cient number of

_plants

for the test. Some flowers from

resistant

_plants

were

_pollinated

with

_Aranguez.

Not more than 20% of these crosses were

_successful,

but

the

_aging

of

_{plants (4-5}

months after

_planting)

led to

an increase in

_{fertility. Only}

13 families

_yielded

proge-nies, generally

5

_plants

_per

_family

were crossed. We

thus obtained 26

_progenies,

named AM 01

_through

AM 26.

_They

were tested for resistance to P solanacearum

in the infected field.

After elimination of the 2 most

_susceptible

families

(AM

02 and AM

_25),

the AM families were crossed in

pairs

at random.

_Twenty-six

families were

obtained,

numbered from SAM 01 to SAM 26.

_They

were

submit-ted to a wilt resistance test in 1986 under artificial

inoc-ulation.

Statistical

_analysis

The Friedman

2-way analysis

of variance

_by

ranks was used

_(Friedman,

_{1937, 1940)}

because of

signifi-cant variance differences within families. The "tests

non

_{paramétriques" procedure}

of the STATITCF

mi-cro-computer program (ITCF, 1988)

was used to test the null

_hypothesis

and to _{compare the families with}

the Kalenda control.

RESULTS

Results for

the 36 FAM

families

are

_presented

in

table

I. A wide

variation

between families was

ob-served,

together

with

a

_{large heterogeneity}

within

families. Out of 36

_families,

20

had

a

resistance

level which

was

_{significantly}

better

than

that of

Kalenda. This

_F

₁

_hybrid,

with

88%

dead

or

wilted

plants provides

evidence of the

_efficiency

and

strength

of the artificial

inoculation.

Concerning

the

AM

families tested

in

the

in-fected field

_{(table II),}

a

_good

level

of

resistance

was

observed, since

19 families

exhibited

< 0%

wilted

_plants.

With 82%

dead

or

wilted

plants,

Kalenda exhibits the

_{highest susceptibility.}

The SAM

families

_generally

showed a very

good

resistance to P

solanacearum;

15 had a

re-sistance level

_{significantly}

better than that of Ka-lenda

_{(table III).}

All the

SAM

families were

char-acterized

_by

a

_good

resistance to fruit anthracnosis

_(data

not

_shown).

With

_regard

to the

_shape

and colour of the

fruit,

this

material has

a _very

wide

_variability.

Fruits are

_long

or

round,

and

_green,

_{pink, purple}

or black in colour. The

_variability

also shows up in the colour of leaves

_(from

green to

_purple)

and flowers

_(from

white to

_purple).

DISCUSSION

The first 2

_generations

after the first

_generations

issued

from the

_{interspecific hybrid}

were

_always

found to be sterile. We noticed that

_aging

of

plants

had a

favorable

effect on

_spontaneous

fer-tility

of

_{interspecific hybrids}

under

_tropical

condi-tions. Similar

results such as

_phenotypic

varia-tion in

the

_offspring

of

_{interspecific hybrids}

have

frequently

been

observed

over the course of time.

The

_Aranguez

_parent

_being

_very

_susceptible

to

P

_{solanacearum,}

the results obtained indicate the

average

dominance of genes

governing

the

resis-tance character in the

selected

FAM families. It is

_impossible

to determine the

_respective

roles of the S

_aethiopicum

and of the P18

_parent

in the

_genetic

control of the resistance in the SAM

_families,

the 2

_original

_parents

_being

resist-ant. The recurrent

selection allowed the

recombi-nation of genes

governing genetic

control of bac-terial wilt resistance from the 2

_parents.

This

genetic

recombination

gave

progenies

with

a

sig-nificantly

better resistance

than that of

Kalenda.

The selection method used here has

_proved

to

be very efficient in such a context where the ge-netic

_variability

is wide and

_probably

_implies

sev-eral

_partly

dominant

_genes.

The

_{polycross phase}

allows the

_expression

of

various

gene

combina-tions,

maintaining

a

_{large proportion}

of the

vari-ability

induced

_{by interspecific}

crosses.

A severe

_screening

test was

_adopted

in

order

to eliminate

_susceptible

and

_weakly

resistant

plants.

This

test

_appears

to be as

effective

as the

natural

infectious

_process,

as

_previously

stated

by

Schmit

₍₁₉₇₈₎

and to

result

in a

better

diffe-rentiation

between resistant

and

_susceptible

plants,

as

_{reported by}

Winstead and Kelman

(1952).

Another

_advantage

is that wilt

_symptoms

were

observed

earlier than

with natural

contami-nation. We believe that the

_validity

of this inocula-tion

_{procedure (mixed}

cultures of bacterial

strains)

for

_breeding

_{programs is}

increased

by

using representative

strains of the

_pathogen

in

term _{of host range}

_{(race classification)}

and bio-chemical characteristics

_(biovar

_{classification).}

This

_strategy

is consistent with other

_eggplant

(Goth

et

al, 1986;

Li et

al,

1988)

and

_potato

(Swanepoel

and

_Young,

₁₉₈₈₎

_breeding

pro-grams.

P solanacearum

strains

_originating

from the French West Indies appear to be

_highly

patho-genic compared

with

strains

from

other countries

(Prior

et al,

1989b),

so

that

the

_genetic

material obtained should have a resistance

suitable

for many countries. This resistance will

be evaluated

in a world - wide

_experimental

_system

_together

with the

_original

_parents

and

with other sources

of resistance.

The

_satisfactory

_shape

and

colour

of

the fruit

of resistant

_genotypes

makes

their

direct

use

possible

after fixation

_by

_{self-pollination.}

The

most resistant

families

can also be used as a

source of resistance to

_{improve existing}

varieties,

or in future

breeding

programs.

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