Proceedings of the XLVII Italian
Society of Agricultural Genetics - SIGA Annual Congress
Verona,
Italy - 24/27 September, 2003
ISBN 88-900622-4-X
Poster
Abstract - 1.38
ARE CLASS B HOMEOTIC GENES INVOLVED IN THE EXPRESSION OF
PARTHENOCARPY IN TOMATO?
I. OLIMPIERI, F.
SILIGATO, R. CACCIA, A. MAZZUCATO
Dipartimento
di Agrobiologia e Agrochimica, Sezione di Genetica, Università degli
Studi della Tuscia, Via S.C. de Lellis snc, 01100 Viterbo, Italy
floral mutants, homeotic genes, parthenocarpy,
tomato
In
flowering plants, the ovary develops into a fruit only after pollination and
fertilization, when the requirements for seed development are thoroughly
fulfilled. The genetic and molecular mechanisms that negatively control ovary
development before pollination are not well understood. A recently proposed
theory postulates that stamens act as regulatory factors towards the fourth
floral whorl, by repressing the development of the ovary until fertilization
has taken place. Accordingly to this theory, the silencing of a class B
homeotic gene has been shown to be the cause of parthenocarpy in apple and a
correlation between stamen disruption and parthenocarpy has been reported in
transgenic tomatoes.
In
tomato, natural or induced parthenocarpic mutants offer a useful method to
regulate fruit production and a suitable experimental system to study ovary and
fruit development. Among them, the parthenocarpic fruit (pat)
mutation deserves particular interest because of its strong expressivity and
the pleiotropic effects that affect anther and ovule phenotype. In such mutant
in fact, the anthers show homeotic transformation into carpel-like structures,
seldom bearing external ovules on the adaxial surface and ovules show arrested
integument growth with absence of meiosis and consequent unviability. To date,
no research has tested the hypothesis that stamen aberrations and parthenocarpy
are driven by cues coming from the altered expression of class B genes in this
genotype.
In
this research, we address this possibility by constructing double mutants
between pat and tomato mutations putatively involved in the B
function, such as stamenless-2 (sl-2, that
shows homeotic conversion of stamens into carpels) and pistillate-2 (pi-2, that
shows strong stamen and subtle petal aberrations). In addition, the expression
patterns of tomato class B (LeAP3 and TM6) and
class B-related (TM8 and TM29) genes have been
studied in time courses of tomato WT and pat mutant ovaries.
F1
hybrids between the pat mutant and sl-2 or pi-2
showed wild-type (WT) phenotype, indicating that the mutations affect different
loci. In F2, whereas pat and sl-2
segregated independently and numerous double mutants could be recovered, for pat and pi-2 no
double mutant could be unambiguously recognized out of 200 progeny plants, thus
suggesting tight linkage. pat and sl-2
showed mainly additive effects in double mutant segregants, while some
phenotypes of the pat mutation were counteracted by a single dose of the sl-2
allele. These results indicated that pat and sl-2
mutants might share only partially overlapping genetic determinants.
RT-PCR based expression analysis revealed that all the four assayed genes were down-regulated in the pat mutant ovary or fruit in comparison with the WT. For those transcripts characterized as being specifically transcribed in ovules (AP3), such lower expression in the mutant could be merely explained by the presence of very few normal ovules inside the pat ovary. However, the down-regulation of LeAP3 might also be directly involved in the parthenocarpic phenotype, because this pattern was also observed in the pat-2 mutant, a different tomato mutation for parthenocarpy that shows normal ovule development. Similarly, the lower expression of TM29 in the pat mutant can be directly related to parthenocarpy, because this gene is normally expressed in the young ovary pericarp.