DISCUSSION
Before TYLCV-DNA probes became available, selection for
TYLCV resistance was based on the absence (or attenuation) of
symptoms in infested fields and in amelioration of yields. Today,
we realize that all tomato cultivars and lines currently available
are, at best, tolerant to TYLCV; although yields and symptoms
have much improved compared with susceptible cultivars, plants
contain various concentrations of virus (12). To the best of our
knowledge, none of the tomato cultivars are resistant (no symptoms,
no virus). The only wild tomato species reported to be resistant
upon whitefly-mediated inoculation was L. chilense LA1969
(21). Mapping and introgression of resistance from L. chilense has
led to a tolerant line, in which virus replication and movement is
slower than in the susceptible near-isogenic line (13). We have
initiated a breeding program with the ultimate goal of obtaining
cultivars totally resistant to whitefly-mediated inoculation. Screening
of accessions of wild tomato species reported to be resistant to
TYLCV has shown that susceptible, tolerant, and resistant individuals
can be found within each accession. This heterogeneity is probably due to the fact that most wild species in their propagation
routine are subject to open pollination, especially if they have
self-incompatibility genes like in L. hirsutum, L. chilense, and L.
peruvianum.
We started our breeding program with a cross between plants
from L. hirsutum accessions LA1777 and LA386 completely resistant
to TYLCV. L. hirsutum LA386 was reported to be resistant
to TYLCV from Jordan; several dominant genes controlled the
resistance (9). L. hirsutum LA1777 was reported to be resistant to
TYLCV from Cyprus (10) and tolerant to TYLCV from Sardinia
and from Senegal and to tomato leaf curl virus from India (8). The
L. hirsutum F1 (LA1777 × LA386) was crossed with L. esculentum,
and a series of selfing were performed. At each generation,
individuals were selected for resistance following repeated, massive,
controlled inoculations with viruliferous whiteflies. Segregation
of progeny for resistance, tolerance, and susceptibility indicated
that resistance and tolerance are controlled by different sets of
genes. Segregation in the BC1F3 population supports a model in
which resistance is controlled by two to three additive recessive
genes. In a given plant, tolerance can be masked by resistance.
These two traits may separate after selfing and segregate at the next
generation. Grafting can also separate resistance and tolerance.
For example, plant number 25 (Table 3) remained symptomless
when grafted on a susceptible infected plant (Table 4, plants C and
D), because it carries a gene for tolerance that is masked by the
genes conferring resistance. Contrary to resistance, tolerance is controlled
by one dominant major gene. The recessive nature of resistance
and the fact that it is a polygenic trait may explain the difficulties
in introgressing this trait into the domesticated tomato.
Grafting experiments have shown that resistance is lost when an
infected susceptible plant is grafted with a resistant plant (as scion
or not): the resistant plant becomes tolerant. Therefore, we suggest
that resistance is expressed at the whitefly-plant interface. Once
the virus is in the vascular system, resistance is lost. Similar results
were reported showing a break of resistance of L. chilense LA1969
and of L. hirsutum LA1777 following TYLCV delivery by agroinoculation
(11). We have now reached a stable BC1F4 line (denominated 902)
that does not segregate for resistance and that has good horticultural
characteristics (80- to 120-g red fruits, fertility, self-compatibility,
vigor, and good plant morphology). This line does not support
virus replication even upon controlled massive whitefly-mediated
inoculation of young seedlings and does not need protection with
nets or insecticides. The F1 with L. esculentum is tolerant, produces
120- to 150-g fruits, and does not need protection from inoculation.