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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
Università degli Studi della Tuscia
Facoltà di Agraria
Dipartimento di Agrobiologia e Agrochimica
Proprietà letteraria riservata
Grafica e lavoro editoriale: Andrea Mazzucato, Mario A. Pagnotta,
Maurizio E. Picarella, Pietro Mosconi
Stampa: Centro Stampa Università della Tuscia
Finito di stampare nel maggio 2008
Con il contributo di: Università degli Studi della Tuscia (Dipartimento
di Agrobiologia e Agrochimica), Società Italiana di Genetica Agraria,
Società di Ortoflorofrutticoltura Italiana (Sezione Orticoltura), Barilla
S.p.A.
DABAC
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
2
TOMATO
GEN omic S
etic
The Italian
Contribution
“Genetica e genomica del pomodoro:
il contributo italiano”
“Tomato genetics and genomics:
the Italian contribution”
Giornata di Studio in occasione del pre-pensionamento del
Prof. Gian Piero Soressi
A workshop organized for the retirement of
Prof. Gian Piero Soressi
29-30 maggio 2008
Facoltà di Agraria, Aula del Consiglio - Università degli Studi della
Tuscia
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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PROGRAMMA
Giovedì 29 maggio
14.30 Registrazione
15.00 saluto del Rettore dell’Università degli Studi della Tuscia
saluto del Preside della Facoltà di Agraria
saluto della Società di Ortoflorofrutticoltura Italiana
saluto della Società Italiana di Genetica Agraria
Il pomodoro: una prospettiva internazionale
Chair: Enrico Porceddu
15.30 Lorenzo Maggioni
BIOVERSITY International
“Il gruppo di lavoro Solanaceae di ECPGR: attivita' di
cooperazione in Europa per le risorse genetiche”
16.00 Fernando Nuez, Jaime Prohens, María José Díez
Università Politecnica Valencia - Spagna
“The Institute for the Conservation and Improvement of
Agrodiversity (COMAV) input to tomato genetics and
breeding”
16.40 Silvana Grandillo
CNR IGV, Napoli
“International SOL Genome Project: il contributo italiano”
17.10 Coffee break
Il pomodoro: una prospettiva italiana I
Chair: Carlo Lorenzoni
17.30 Agostino Falavigna
CRA ISPORT Montanaso Lombardo
“Contributo di Gian Piero Soressi all'avvio ed alla crescita
scientifica a Montanaso Lombardo”
17.50 Andrea Mazzucato
Università della Tuscia
“Il Codice ‘Soressi’ ”
18.10 Alberto Pardossi
Università di Pisa
“Incontro tra breeder e agronomi: il caso delle colture
idroponiche”
18.30 Roberto Ranieri, Marco Silvestri
Barilla S.p.A.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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“L'esperienza Barilla nella selezione del pomodoro da industria”
20.30 Cena sociale
Venerdì 30 maggio
Il pomodoro: una prospettiva italiana II
Chair: Mario Pagnotta
9.00 Luigi Ricciardi, Concetta Lotti
Università di BARI
9.30
“Il gene ol-2 di resistenza all'oidio”
Gian Piero Soressi
Università della Tuscia
“Mutagenesi classica e variabilità somaclonale in pomodoro:
metodologie sperimentali a confronto e loro sinergie”
10.00 Pierdomenico Perata
Scuola Sup.re Sant’Anna, Pisa
“Produzione di antociani in Arabidopsis e pomodoro”
10.30 Carlo Rosati
ENEA Casaccia, Roma
“Carotenoids: key players in tomato fruit ripening”
11.00 Coffee break
Il pomodoro: una prospettiva italiana III
11.20 Francesco Cellini, Filomena Carriero
METAPONTO Agrobios
“Uso di mutanti per studi di genetica funzionale in pomodoro:
l’esperienza di Agrobios dal Blind al TILLING”
11.50 Luigi Monti
Università Federico II, Napoli
“Il progetto GenoPOM”
12.20 Francesco Salamini
Università di Milano
“Conclusioni e prospettive”
12.50 Gian Piero Soressi
“Chiusura dei lavori”
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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ABSTRACTS
“The Institute for the Conservation and Improvement of Agrodiversity
(COMAV) input to tomato genetics and breeding”
Nuez, F.; Prohens, J.; Díez, M.J.
Instituto de Conservación y Mejora de la Agrodiversidad Valenciana (COMAV).
Universidad Politécnica de Valencia, Valencia, Spain. [email protected].
The tomato is the most economically important vegetable crop in Spain. The
Institute for the Conservation and Improvement of Valentian Agrodiversity
(COMAV) leads the Spanish tomato breeding in the following research areas:
genetic resources, breeding for resistance to diseases, breeding for quality,
parthenocarpic fruit set. The COMAV holds a collection of almost 4000
accessions of tomato and wild relatives mainly from Spain and LatinAmerica,
but also from many other countries. Intensive studies are being conducted on
the morphological and molecular diversity of both the cultivated tomato and its
wild relatives. Research on resistance to viral diseases is being conducted with
Tomato yellow leaf curl virus, Tomato spotted wilt virus, Tomato mosaic virus,
Pepino mosaic virus, and Parietaria mosaic virus. Development of resistant
cultivars and research on diagnostic methods and molecular aspects is being
carried out. Studies on organoleptic and nutritional quality have focused on the
search of sources with high content of sugars, acids and antioxidants, and in the
development of rapid and accurate methods to measure the content of these
constituents. The environmental effect and the genetic control of some of these
components have been elucidated. Genetic control of parthenocarpy, as well as
physiological and molecular aspects have been also studied. We are comparing
the transcriptome of parthenocarpic and non-parthenocarpic fruit set in several
tomato lines to identify the main pathways implicated in this process. A
strategy to identify the genes responsible for parthenocarpy in these lines has
been started using a positional cloning and candidate genes approaches.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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“Contribution of Gian Piero Soressi to scientific grow of the Montanaso
Lombardo Research Section”
Agostino Falavigna
CRA- Unità di Ricerca per l’Orticoltura di Montanaso Lombardo (LO)
Gian Piero Soressi started his scientific activity at the Montanaso Lombardo
Section of the Research Institute for Vegetable Crops of Pontecagnano (SA) on
May 1968 as a researcher. Since 1973 till October 1983 he was Director of the
Section.
During this period he started breeding programs on tomato, asparagus, onion,
bean and an agronomic project on the utilization in agriculture of waste heat
from electric power stations.
Gian Piero Soressi addressed each one of his young collaborators to specialize
on a single crop and spent a big care to their scientific growth. Although not
very common at that time and for the Research Centres of the Ministry of
Agriculture, he pushed the collaborators to spend a period abroad in order to be
trained in outstanding International Research Institutions.
Modern plant breeding may help food branded companies in the daily fight in
the market because the new tools offered by genomics allow a very efficient
selection of the traits required for some end product innovations.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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“The Soressi’s code”
Andrea Mazzucato
Dipartimento di Agrobiologia e Agrochimica, Università degli Studi della Tuscia, via
S.C. de Lellis, s.n.c., 01100 Viterbo (Italy) – [email protected]
The Soressi’s code does not refer to the genetic code, as would be expected in a
workshop of geneticists, nor to professional or ethical codes, as would be
expected in celebrating the career of a honest and enthusiastic worker. More
simply, we refer to the code that every geneticist has to give to his materials,
accessions and generations.
Soressi is first a ‘code maker’: since the beginning of his career in the early
60s’ he obtained by mutagenesis a considerable number of mutants that were
coded, named, multiplied, described and studied in his research activity. Beside
genetic codes, Soressi invented the phenotypic codes, that are a list of symbols
used in labels to phenotype more effectively large collections in field trials.
Second, Soressi is a ‘code collector’: he did one of the first collections of
tomato landraces some forty years ago, when the ‘hybrid era’ was not yet
commenced, without forgetting to single out spontaneous mutants segreganting
in swards. Then, he collected materials from colleagues having the most
important germplasm accessions in the tomato breeding history, as the
collection of ‘color genes’ from L. Butler and L.A. Darby, the mutant
collections of H. Stubbe and C.R. Rick, the near isogenic lines from J.
Philouze, the introgression lines selected by D. Zamir.
Beside collecting, Soressi has been studying all his entries as a ‘code user’. For
many mutations he deeply studied the expressivity, allelism and potentialities in
breeding. In some experiments, he put together sets of dozens of mutations, as a
prodrome of the modern ‘-omics’ approach.
Finally, he has been and is a ‘code disseminator’, as he distributed the materials
he characterized to the scientific community, together with the information and
many times with related ideas and suggestions. This sowing of the Soressi’s
code has been really extensive and is germinated in many of the advances on
tomato genetics and genomics nowadays.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
8
“A joint venture between breeders and agronomist:
the case of soilless culture”
Alberto Pardossi
Dipartimento di Biologia delle Piante Agrarie – Università di Pisa
Viale delle Piagge, 23 – 56124 Pisa
[email protected]
The presentation illustrates the long cooperation between Prof. G.P. Soressi
(University of Tuscia) and Prof. A. Pardossi (University of Pisa), which was
initiated more than 20 years ago.
At first, the work focused on the development of new tomato genotypes
suitable for greenhouse soilless culture, in particular for high density ((10-14
plant/m2), short-cycle (1-2 trusses per plant) cropping systems to be carried out
without the use of auxin treatment for fruit set. With this goal, a series of
experiments were conducted between 1987 and 1992 to investigate the
adaptation of different monostem (to-2) and parthenocarpic (pat-2) genotypes
to NFT (nutrient film technique). Three main types were identified for their
vegetative and reproductive characteristics. It was also found that the
monostem genotypes developed normal axillary shoots when grown with high
nitrogen content in the nutrient solution.
More recently, the cooperation between Proff. Soressi and Pardossi was
extended to Prof. R. Izzo research team (also at University of Pisa) to
investigate the influence of salinity on fruit qualità of tomato plants grown in
closed-loop rockwool culture. In this study, conducted in the framework of
PRIN projects, three near-isogenic lines of cv. Gimar differing, with respect to
wild-type, in ethylene production and fruit ripening pattern were used: gf (green
flesh), nor2 (non-ripening2) and Nr (Never ripe). The same response was
observed in all genotypes: the higher salinity growth solution increased dry
matter content, total soluble solids (TSS), titratable acidity and ascorbic acid,
but reduced crop yield. Our data confirm the lack of any relationship between
the larger accumulation of TSS and ethylene production in the fruits of salttreated plants. Salinity induced an oxidative stress, and the sensitivity to salt
treatment was genotype-dependent; for instance, Gimar gf was much less
sensitive.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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“Barilla Experience in Processing Tomato Selection”
Roberto Ranieri e Marco Silvestri
Barilla G. e R. Fratelli – Via Mantova 166 – 43100 Parma
Corresponding author: [email protected]
The main areas where branded companies are active to deliver innovative end
products are: technology/processing, service/packaging, function/usages and
raw materials, which represent a powerful tool of differentiation.
Barilla’s approach on a strategic raw material is to know in depth the whole
production chain and to understand the critical points to study and to develop
research projects. Often, the answer is breeding. Conventional and molecular
assisted breeding. Adopting breeding on durum wheat (Triticum turgidum var.
durum Desf.) and on processing tomatoes (Solanum lycopersicum L.), Barilla
has been able to differentiate some of its products in terms of cost, texture,
nutrition and appearance.
For tomatoes, a good example is the “Scarpariello case”, a tailor made tomato
variety selected by Barilla and Peotec Seed company and used by Barilla to
produce outstanding and challenging branded sauces.
Industrial sauces business has great potentialities to further develop since, in
Italy, for less than 10 % of the pasta dishes industrial sauces are used but
usually sauces are still home made. People are still not much confident in
adopting industrial sauces because the common idea is that “sauces are all the
same”, “all sauces have industrial taste” and “ingredients are not natural”.
Sauces containing “Scarpariello” are a challenge to change this perspective.
“Scarpariello” variety was selected for outstanding and taste which withstands
during the thermal applications to guarantee the best raw material quality. It
allows obtaining innovative and distinctive sauces that have been “branded”
with the variety name too. In the future, through breeding, it could be possible
to differentiate even more among sauces in terms of taste and why not, color
and nutritional traits too.
The raw material relevance on final product innovation is so distinctive that
“Scarpariello story” become the core of the TV advertising campaigns of both
pasta and sauces categories.
Modern plant breeding may help food branded companies in the daily fight in
the market because the new tools offered by genomics allow a very efficient
selection of the traits required for some end product innovations.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
10
“The tomato ol-2 gene confers resistance to Oidium neolycopersici”
Ricciardi L.*, Lotti C.**,Pavan S.*, Visser R.***, Bai Y.***
*) Department of Agroforestry, Enviromental Biology and Chemistry, Genetics and Plant Breeding
Unit, University of Bari, Bari 70126, Italy
**) Department of Agro-Environmental Science, Chemistry and Crop Protection, University of
Foggia, Via Napoli 25, Foggia 71100, Italy
***) Laboratory of Plant Breeding, Plant Sciences Group, Wageningen University, 6700 AJ
Wageningen, The Netherlands
The tomato recessive gene ol-2 (conferring resistance to Oidium
neolycopersici) was identified at University of Bari in an accession of Solanum
lycopersicum L. var. cerasiforme and localized on chromosome 4 by means of
AFLP, RAPD and SCAR markers, using a F2 population obtained crossing the
resistant accession, named LC-95 and the susceptible cultivar SuperMarmande.
Further studies were carried out in co-operation with Wageningen University in
order to study the resistance mechanism of ol-2, observing that ol-2 confers
papilla-associated and race-non-specific resistance to tomato powdery mildew
caused by O. neolycopersici. Based on homology with recessive genes
conferring resistance to powdery mildew in other species, such us barley and
Arabidopsis, the tomato candidate gene (named LeMLO1) was identified in
Solgenes data base and the comparison of nucleotide sequences of resistant and
susceptible genotypes revealed a 19 bp deletion in the resistant parent. In the F2
of a cross between a resistant line bearing ol-2 and the susceptible tomato
cultivar (cv.) Moneymaker, a chromosomal fragment disrupting the LeMLO1
coding region co-segregated with resistance. This polymorphism results in a
frame shift and thus a truncated non-functional LeMLO1 protein. As both the
genetic and the phytopathological characteristics of the ol-2-mediated
resistance are reminiscent of powdery mildew immunity conferred by loss-offunction of mlo (mildew resistance locus o) alleles in barley and Arabidopsis,
we initiated a candidate gene approach to clone Ol-2. The LeMLO1 was
mapped in the chromosomal region harbouring the Ol-2 locus by using both
genetic and cytogenetic tools. Complementation experiments, using transgenic
tomato lines and virus-induced gene silencing (VIGS) assays, suggested that
loss of LeMLO1 function is responsible for powdery mildew resistance
conferred by ol-2. Our researches reveal the second example of a natural mlo
mutant, suggesting that natural mlo polymorphisms may compensate for the
man-made poverty of resistance gene diversity in the gene pool of cultivated
species.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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“Classical mutagenesis and somaclonal variation in tomato: confronted
procedures and synergies”
Gian Piero Soressi
Dipartimento di Agrobiologia e Agrochimica, Università degli Studi della Tuscia, via S.C. de
Lellis, s.n.c., 01100 Viterbo (Italy) – [email protected]
This presentation aims, at the same time, to be a sign of gratitude and warning:
a deep thanks to my first scientific leaders, proff. Angelo Bianchi and Gian
Tommaso Scarascia Mugnozza, scientists in radiogenetics respectively for
maize and wheat; a warning for the young researchers under way of tomato
genetics and breeding. What I have learnt since the preparation of my
graduation thesis on the effects of x-rays on tomato seeds, were the terms
“gametic” and “intrasomatic selection”, “true control” and the symbols
indicating the different plant progenies issued from the non treated, X0, X0S1,
and x-rayed, X1, X2, seeds. Remembering that X1 is analogous to F1, I was
surprised in realizing that in most published papers on in vitro induced
somaclonal variation, R0 took place of R1 (like X1), the segregating progeny
became R1 (instead of R2, like X2), therefore the true R0 like X0 disappeared.
Such an inconsistent change, also disapproved by prof. Francesco D’Amato, led
to underestimate the importance of the “true control”, so that some pre-existing
mutation of the seeds used for obtaining the explant were de facto
misinterpreted as somaclones. Unfortunately (or fortunately), I saw with my
own eyes some of such mistakes when I was asked to check a few very
interesting, putative somaclones. With such a store of experience, when later
we also decided to evaluate the efficiency and quality of the in vitro inducible
variation, we took care of having a “true control”, by setting up a procedure
that permit the recovery of the explant donor seedling. The same procedure was
applied in the experiments where we combined chemical with in vitro
mutagenesis. In this way we were able to distinguish and separately screen all
the plant progenies under study: M0R0 [from non-treated (M0) and nonregenerated (R0) seedlings]; R0M1 (from EMS treated seeds); R1M0 (from
cotyledon explant regeneration only); R1M1 (regenerated plantlets from
cotyledons of EMS-treated seeds). Through this research we succeeded in
obtaining somaclonal variants induced by chemical mutagenesis but recovered
via cotyledon regeneration. A proof of this event was an anthocyaninless
mutant singled out in a R1M1 progeny. This mutant likely induced by EMS in a
cotyledon cell, should have been lost without cotyledon regeneration because it
was induced in a tissue that do not give contribution to sexual lineages. No
anthocyaninless somaclone has been up to now reported as a result of the sole
in vitro cotyledon regeneration.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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“Arabidopsis thaliana MYB75/PAP1 Transcription Factor induces
Anthocyanin Production in Transgenic Tomato Plants”
Pierdomenico Perata
Scuola Superiore S. Anna, Pisa
Tomato (Solanum lycopersicum L.) cv. Micro-Tom plants were transformed
with the Arabidopsis thaliana (L.) Heyhn MYB75/PAP1 (PRODUCTION OF
ANTHOCYANIN PIGMENT 1) gene. This gene codifies a transcription factor
which is involved in anthocyanin production and is modulated by light and
sucrose. The transgenic tomato plants expressing AtMYB75 were characterized
by a significantly higher anthocyanin production under normal growth
conditions in leaves, stems, roots, flowers and, interestingly, in fruits.
Anthocyanin accumulation was not widespread but took place in specific
groups of cells located in epidermal or cortical regions, or in proximity to
vascular bundles. In the vegetative organs of the transgenic plants, where
AtMYB75 overexpression was determined, a clear up-regulation of all the main
genes involved in flavonoid pathway was also detected. On the contrary, no
effect was produced on the expression of the tomato MYB-gene ANT1
(ANTHOCYANIN1) that had previously been identified as a transcriptional
regulator of anthocyanin biosynthesis. Additionally, induction of many but not
all the structural genes of the biosynthetic pathway was observed in the fruits.
The higher basal content of anthocyanins in the leaves of the transgenic plants
could be further increased in the presence of high light conditions and
contributed to mitigate photobleaching damages under high irradiance.
Transformations of Anthocyanin fruit (Aft) and Ailsa Craig tomato genotypes
were also performed obtaining similar results. Molecular characterizations of
these transgenic plants are in progress.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
13
“Carotenoids: key players in tomato fruit ripening”
Carlo Rosati1, Gianfranco Diretto2, Raffaela Tavazza2, Florence
Bouvier3, Bilal Camara3, Einat Bar4, Efraim Lewinsohn4, Nicholas
Schauer5, Alisdair Fernie5, Giovanni Giuliano2
1
ENEA Trisaia, S.S.106 km 419+500, 75026 Rotondella (MT)
ENEA Casaccia, via Anguillarese, 00060 S. Maria di Galeria (RM)
3
CNRS, Plant Molecular Biology Institute, 12 rue du Général Zimmer, 67084 Strasbourg, Francia
4
Institute of Plant Sciences, Newe Ya’ar Research Center, Agricultural Research Organization,
P.O. Box 1021, Ramat Yishay, 30095 Israele
5
Max-Planck Institute for Molecular Plant Physiology, Am Muehlenberg 1, 14476 Golm,
Germania
2
Carotenoids are C40 plastid-associated molecules which impart
nutritional properties and visual appeal to tomato. Due to the strong
accumulation of carotenoids in the fruit and the availability of mutants,
tomato has long been used as a model species for the carotenoid
pathway. The biosynthetic route has been extensively characterized also
by metabolic engineering, which have revealed some critical regulatory
steps of the carotenoid content in tomato fruits and successfully
increased pro-vitamin A and xanthophyll content. Comprehensive and
targeted gene expression and metabolite analyses of tomato
transformants showed that carotenoid cyclization/hydroxylation is also
able to induce global changes in fruit ripening, by modulating fruit
transcript levels and metabolic profiles, thereby affecting important
traits such as fruit firmness, ABA levels, and the emission of ethylene
and volatile compounds.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
14
“Uso di mutanti per studi di genetica funzionale in pomodoro:
esperienza di Agrobios dal Blind al TILLING”
Francesco Cellini, Filomena Carriero
Metapontum Agrobios, S.S. Jonica 106, km 448.2, 75010 Metaponto (MT) Italy
La disponibilità di mutanti è di straordinaria importanza per il
miglioramento genetico di colture in ambito agrario e per la ricerca della
funzione di geni di interesse. Le tecnologie per la generazione e lo
studio di mutanti hanno registrato notevoli sviluppi, grazie al progresso
delle tecniche molecolari ed alle conoscenze acquisite attraverso il
sequenziamento del genoma delle piante superiori.
La disponibilità di particolari strumenti molecolari ha consentito di
mettere a punto, nel corso degli anni, sofisticati approcci di genetica
funzionale, quali il transposon tagging eterologo, il gene walking che
hanno evidenziato successi e fallimenti in modo variabile.
Un esempio dell’evoluzione di tali tecnologie è fornito dall’approccio
seguito da Agrobios per l’isolamento e la caratterizzazione del gene
blind, e della sua variante allelica torosa,in pomodoro, un caso che
dimostra l’importanza di disporre di mutanti ben caratterizzati.
Negli ultimi anni è apparsa una nuova potente tecnologia che consente
di identificare nuove varianti alleliche di geni, la cui sequenza
nucleotidica è nota, e di ottenere nuovi genotipi migliorati nel carattere
di interesse. Questa tecnologia, ad oggi applicata su diverse specie, è
conosciuta con il nome di TILLING (Targeting Induced Local Lesions
In Genomes).
Precisamente il TILLING utilizza la mutagenesi chimica (es. EMS) per
indurre variabilità (mutazioni) nel genoma di un organismo e tecniche di
biologia molecolare (PCR) per individuare mutazioni utili e quindi
varianti alleliche di geni di interesse. Attraverso questa tecnologia
possono essere selezionati genotipi con nuove caratteristiche
agronomico-nutrizionali direttamente trasferibili verso uno sviluppo
commerciale.
Presso la Metapontum Agrobios, l’applicazione della tecnologia
TILLING ha avuto inizio con la produzione, mediante mutagenesi
chimica (EMS), di una collezione di mutanti di pomodoro (cv Red
Setter) costituita da 6677 Famiglie M2 e 5508 Famiglie M3.
Dati sullo sviluppo della piattaforma TILLING relativi sia alla
produzione della collezione di mutanti di pomodoro che agli screening
molecolari verranno presentati e discussi.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
15
SINOPSI DELL’ATTIVITÀ SCIENTIFICA DEL PROF.
GIAN PIERO SORESSI
L’attività scientifica di Gian Piero Soressi, pur mantenendo il comune
denominatore dell’utilizzo della variabilità naturale e indotta per il
miglioramento genetico di specie orticole (il pomodoro innanzitutto), ha
affrontato un gran numero di argomenti, sotto diverse prospettive e,
soprattutto, con l’interazione di tantissimi studenti, collaboratori e
colleghi. Crediamo che questo prezioso lavoro, a volte rimasto al di
fuori della letteratura internazionale più diffusa, meriti una sintesi più
puntuale rispetto a una semplice lista di pubblicazioni.
SYNOPSIS OF PROF. GIAN PIERO SORESSI’S
SCIENTIFIC ACTIVITY
The research activity of Gian Piero Soressi, although maintaining the
common denominator of using natural and induced variation for the
genetic improvement of vegetable (mainly tomato) crops, has addressed
a myriad of research topics, under a number of perspectives and, most
notably, interacting with a great number of students, collaborators and
colleagues. We believe that this valuable work, sometimes remained
outside of the most diffused international literature, deserves a
presentation somehow more detailed than a simple list of publications.
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
16
7B-1 mutant
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
The conditional male sterile mutant 7B-1, showing sterility in LD and
fertility in SD, has been the object of a
collaborative research with Prof. V.K.
Sawhney of the University of Saskatoon,
Canada. The expressivity of the mutation
was characterized in the Italian conditions
at Viterbo and sterility was very good in
spring-summer (see Figure). However, the
recovering of fertility in our autumn
Sterile phenotype of 7B-1 in
conditions was not satisfying (10 seeds per
spring LD conditions (left)
fruit on average). Creating artificial SD in
and partial recovering of
the spring season by covering the plants
fertility in SD in autumn
was proposed as a promising alternative. A
(right).
possible allelism between 7B-1 and the
class B MADS-box gene DEFICIENS was excluded by mapping
experiments (Stamegna 2008).
Aloe arborescens
Chiara Bedini, Antonio Tiezzi (Università della Tuscia, Viterbo Italy)
A micropropagation protocol for Aloe arborescens has been developed in
order to maximize the multiplication index and to minimize the cycle
length. Explants were sterilized in NaOCl and subcultured weekly to
overcome the effects of released polyphenol
that, otherwise, caused browning of the cut
surface, tissue damage and death. Five
substrates having Murashige & Skoog salts as
a basis and differing in the type and
concentration of auxins and cytokinins were
tested. Explants cultured on the substrate
containing 1.0 mg l-1 NAA and 2.0 mg l-1 BA
showed abundant sprouting from their
Aloe explant with an
basement and from axillary meristems, with a
axillary sprout obtained
mean multiplication index of 3.5. Biological
in vitro.
effects of leaf extracts from micropropagated
plants of A. arborescens on proliferation of the murine myeloma cell line
P3X were evaluated. Preliminary results confirmed the presence of
bioactive molecules and their activity in inhibiting cell proliferation; such
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activity was lower than that of analogous leaf extracts from three-year old
plants (Bedini et al. submitted).
Anthocyanins
Mariagrazia Antonelli, Andrea Mazzucato, Enrico Santangelo (Università della
Tuscia, Viterbo Italy)
Anthocyanin pigments belong to the class of phenolic compounds named
flavonoids and are biosynthesized via the shikimic acid pathway. They
provide for red, blu or purple colour in flowers, fruits, stems, roots and
leaves, playing an important role in plant reproduction by recruiting
pollinators and seed dispersors. Anthocyanins in plants act also in
protection against various stresses such as UV irradiation, hight light
intensity, ROS, insect and pathogen attacks. In the last years, interest in
anthocyanins has intensified because of their health benefits as dietary
antioxidants and their pharmacological properties for therapeutic purposes.
In tomato, flavonoids are accumulated in a tissue-specific way depending
a
b
sc
Tomato plant originated from the cross atv x Aft and Aft x atv : a) “sun
black” with high anthocyanin fruit pigmentation; b) selected plant suitable
for mechanical harvesting with typical Aft pigmentation.
on the fruit developmental stage. They are mainly synthesized in the
epicarp where naringenin, a precursor in anthocyanin biosynthesis, is the
main component reaching a peak at the fruit turning stage. Soressi’s
collection of tomato germplasm contains accessions of spontaneous or
artificially induced anthocyanin mutants, obtained since the 60s, and
genetic stocks from the C.M. Rick Tomato Genetics Resource Center. This
collection includes monomendelian mutants characterized by a remarkable
increment (ag, atv, atvc, atvsc, Aft, Abg), decrement (al) or absence (a, aa,
asc, ac, af, ah, aw, bls) of anthocyanins in stem, leaf and fruit.
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Using spectrophotometric analysis, Santangelo et al. (2002), studied the
modifications occuring in tomato fruit pigments during ripening. They
found that in the Aft mutant, anthocyanin content decreased during fruit
ripening as for chlorophyll. Subsequently, by using fluorescent nondestructive techniques, Lai et al. (2007) analized changes in anthocyanins,
chlorophyll, carotenoids in epicarp of tomato during fruit ripening,
identifying the main fluorescent bands of these pigments. These results
could give important information about the optimal conditions for fruit
harvesting using non-destructive techniques.
Considering the biological relationship between anthocyanin induction and
insect attack, experiments have been carried out in order to verify the effect
of tomato leaves with different anthocyanin content (atvsc, a and the control
UC82) in the diet of Spodoptera littoralis. Results suggested that high
anthocyanin content could give sub-letal effects to the physiology of the
larvae (see Spodoptera littoralis).
TOMANTHO, a MIUR-National Project (2006), coordinated by Prof. P.
Perata from the Sant'Anna School for Advanced Studies of Pisa, involving
Soressi’s group as partner, aims to investigate different aspects connected
with anthocyanin content in tomato leaves and fruits by using atv, atvc, atvsc
and Aft mutants. Objectives of this project are: (i) development of tomato
breeding lines with high anthocyanin content in leaf and fruit, (ii)
identification of molecular markers linked to the genes atv and Aft, (iii)
evaluation of the response of tomato anthocyanin mutants to the
phytopathogenic baterium Xanthomonas vesicatoria, (iv) study of
phenotipic and agronomic interaction between anthocyanin genes and
pigment enhancer genes (see Tomato, “high-pigment genes”). Today, F4
and F5 progenies derived from the atv x Aft and Aft x atvsc crosses have
been obtained, and the other objectives are being realized. Within the F4
progenies some double recessive recombinants named “sun black” for their
unusual fruit pigmentation, were singled out (see Figure).
A previous MIUR-National Project (2005) in collaboration with Prof. M.
Badiani at the “Mediterranea” University of Reggio Calabria on the
response of some tomato mutants to ozone exposure (see Ozone) showing
that the atv mutant seems to be more resistant to O3 than other tomato
genotypes, confirming a role of anthocyanins as ROS scavengers.
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Artichoke
Enrico Santangelo (Università della Tuscia, Viterbo Italy)
In the 70’s, through multivariate analysis of 40 relevant phenotypic traits of
the artichoke traditional varieties cultivated in Italy, four major affinity
groups namely “Spinosi”, “Catanesi”, “Violetti”, “Romaneschi” were
recognized. In the framework of the breeding activity developed in our
country by R. Tesi and A. Graifenberg from the beginning of the 80’s and
relatively more recently by F. Saccardo’s team, rightly enters the
enthusiastic, 15 years-selection work of Pietro Papalini. Soressi and
Papalini met for the first time in the 60’s for their contribution in trying to
solve the problem of the mechanical harvesting of the tomato crop in Italy.
The selection work on artichoke by Papalini has been mainly based on
open-hand crosses between
clones of different cultivars
from the “Romanesco” and
“Violetto”
groups.
By
applying such a methodology,
he selected a number of
clones well characterized for
plant vigour, yielding period,
shape, size, commercial and
organoleptic quality of the
heads. To continue his work
Plant of the cv Giove showing the nearly
of open-hand crossing and
homogeneous ripening of 1st and 2nd order
selection, Papalini created a
heads.
basic “genetic pool” named
by him “Club della Biodiversità” formed by his best selected clones
(Gigante, Moro di Corneto, Pacific, Marengo, Chiarostar, Etrusco, Apollo,
Exploter, Roma99), integrated afterwards with new entries endowed with
special traits required by fresh market or industry. At this point Papalini’s
experience met the Soressi’s one as geneticist and breeder (Soressi 2003).
From this cooperation, research on artichoke genetics continued in the
frame of the experimental PRAL program “Artichoke Varieties Innovation
and Cultivation Planning” supported by the Lazio Region and coordinated
by the phytopathologist Prof. Paolo Magro. The activity carried out in two
years of research included the selection of new F1-open clones and a bioagronomic field trial in which the traditional cv. Castellammare, new clones
of Romanesco type (Grato1, Grato2, C3), cv. Apollo (Vitroplant), Exploter
and Giove (both from Papalini) were compared. This field trial proved the
earliness of C3 compared with the other clones. Each clone showed distinct
plant habit and head features (size, shape and colour nuance). Apollo
exhibited a peculiar minor leaf insertion angle possibly responsible of its
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higher tolerance to cold and to other unfavourable climatic events. Giove
reached the maximum concentration of head emission permitting to harvest
all the 1st and 2nd rank heads only 9 days after the main head (see Figure);
thus it appeared particularly suitable for artichoke production destined
either to fresh market or to the industry. The selection activity carried out
on artichoke appears promising even considering the difficulties in putting
together in few clones all the desired traits. The “gene-pool” established
with the various types of the artichoke clones enriched by the best F1-open
clones selected by Papalini in 2005 is a valid premise for selecting
promising F1-clone combinations among the new F1-open plants from the
open-hand crosses obtained in 2006 and observed during the 2008, first
year of growth.
Asparagus
Agostino Falavigna (CRA – Research Unit for Vegetable Crops, Montanaso
Lombardo, Lodi Italy)
In Italy the unique breeding program on asparagus, promoted and started in
1974 by Soressi, at that time head of the ISPORT Section (Soressi and
Falavigna 1979; Falavigna and Soressi 1981a), is still working at CRA –
Research Unit for Vegetable Crops of Montanaso Lombardo. Such a
program has been constantly financed by the Emilia Romagna Region
(through CRPV in Cesena), Veneto Region (through Veneto Agricoltura)
and the Italian Ministry of Agriculture in the sub-project “Asparagus and
Onion” (Soressi 1986b; 1988).
Since the conventional breeding methods based on selfing cannot be easily
applied to asparagus, a dioecious and perennial species, the research
activity progressed through the “Diego” F1 hybrid (Falavigna et al. 1982,
XXVI SIGA Annual Congress), obtained by crossing two heterozygous
plants (selected few years for their spears yield and quality), until to the
double haploid (DH) clones; in both cases the parent clones were
micropropagated (Falavigna and Soressi 1981b; Falavigna et al. 1984a).
To obtain DH clones, in vitro anther culture was sistematically applied (Fig.
1, 2). The “super-male” androgenetic DH clones originate all-male hybrids
(AMH) of two kinds: two-ways AMH, when both parents are DH clones;
three-ways AMH, when the seed parent is an heterozygous clone.
All DH clones were evaluated in fields naturally infested by Fusarium spp.
(the most important soil born disease of asparagus) and selected for plant
survival ratio, fertility and morphological traits.
During thirty years of work, about 300,000 anthers of Italian asparagus
landraces and hybrids from abroad were cultured. The in vitro culture
technique on average yielded 1 androgenetic clone out of 100 anthers and
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the percentage varied from 0.1 to 30 according to the anther donor
genotype. Out of 3,000 androgenetic clones obtained, 58% were DH and
the female/male ratio was near 1/1. The selection pressure over the DH
clones was about 95%; therefore 103 and 100 respectively female and male
clones were selected and systematically utilized as parents of all male F1
hybrids (Falavigna et al. 1983; 1984b).
Fig.1 Callus proliferating
from a pollen grain in
asparagus anther.
Fig.2 Androgenetic calli regenerating
asparagus
stems
ready
for
micropropagation.
Seedling growth analysis was
carried out by comparing 3 new
AMHs with commercial hybrids and local populations (Stancanelli 1989;
Stancanelli et al. 1990). Moreover, a collection of asparagus wild species
germplasm in Sicily, previous botanical classification, was tested for seed
germination and in vitro vegetative propagation (Fig. 3; Venezia 1991;
Venezia et al. 1993). Out of 350 F1 all male hybrids tested so far, ten have
been included in the National Variety Catalogue and four (Eros, Marte,
Ercole, Zeno, Fig. 4) are grown at present on almost the whole green
asparagus
cultivation
area and on a great part
of that cultivated for
white spear production
in Northern Italy (about
2,000 ha). Besides, the
AMH Italo is risen in
Southern
regions
(Falavigna 2007).
Further improvements
Fig.3
Asparagus
for spears yield and
Fig.4 A plant of the
spears of the two
wild
species
A.
quality, plant disease
most important F1
maritimus.
all-male hybrids.
resistance
and
salt
tolerance are expected
on the basis of multifaceted investigations under way by the Falavigna
research group (Falavigna et al. 2008).
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Association genetics
Andrea Mazzucato, Maurizio Enea Picarella (Università della Tuscia, Viterbo Italy)
The characterisation of very diversified materials with molecular markers
offers a unique opportunity to define significant marker-trait associations of
biological and agronomic interest. In this research, carried out in
collaborations with colleagues of the university of Ancona (prof. Papa and
Nanni) and Perugia (prof. Negri and Veronesi), 50 tomato landraces
(mainly collected in central Italy) nine vintage and modern cultivars, and
two wild outgroups were characterised for 15 morpho-physiological traits
and 29 simple sequence repeat (SSR) loci. The markers were selected to
include a group of loci in regions harbouring reported quantitative trait loci
(QTLs) that affect fruit size and/or shape (Q-SSRs) and a group of markers
that have not been mapped or shown to have a priori known linkage (NQSSRs). Compared to the low molecular polymorphism reported in tomato
modern cultivars, our data reveal a high level of molecular diversity in
landraces. Such diversity has allowed the inference of the existence of a
genetic structure that was factored into the association analysis. As the
proportion of significant associations is higher between the Q-SSR subset
of markers and the subset of traits related to fruit size and shape than for all
of the other combinations, we concluded that this approach is valid for
establishing true-positive marker-trait relationships in tomato (Mazzucato et
al. 2008a).
atroviolaceum (atv)
see Anthocyanins
Auxin
see Reporter gene; parthenocarpic fruit (pat); Tomato, leaf
peroxidases
Bean, biotechnological approach
Emilio Mendoza de Gyves, Enrico Santangelo (Università della Tuscia, Viterbo
Italy)
Andrea Allavena (Res. Unit for Floriculture and Ornamental Species, Sanremo
Italy)
Ali Akbar Habashy (Agricultural Biotechnology Research Institute, Karaj Iran)
Prof. Soressi’s activity on in vitro tissue culture, somatic embryogenesis
and genetic transformation of bean (Phaseolus spp.) begun in the 80’s at the
Research Institute for Vegetable Crops of Montanaso Lombardo (Lodi)
(Castelnuovo 1984; Angelini Rota 1987). During the 90’s the efforts
continued in the “In vitro Culture Laboratory” at DABAC (Tuscia
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University). At the end of the 90’s, the group participated to an European
Concerted Action (see Phaselieu Concerted Action) and to the Phaseolus
Italian Network (PIN).
Studies were conducted to determine the effect of growth regulators on the
micropropagation aptitude of several genotypes of common (P. vulgaris)
and running (P. coccineus) bean coming from different countries (Mexico,
Peru, Iran, Italy, Colombia, etc.). Axillary and apical bud explants were
cultured to ascertain any different response to in vitro culture conditions.
Multiple buds production and shoot elongation, with different level of
response, were obtained from all genotypes: BAP was able to give a
satisfactory percentage of multiplication, but, at all concentrations used, it
induced abundant callus at the base of the explants therefore reducing
adventitious bud formation in all cultivars (Allavena et al. 1994; Mendoza
de Gyves 1995).
A protocol for efficient plant propagation via axillary bud explants was
proposed for cv. Montecarlo and with some adjustments was successfully
utilized for other genotypes such as Giulia, Pinto, Clio, etc., including the
running bean, which is more amenable to tissue culture than common bean
(Mendoza de Gyves 1995).
Mature seeds of the Italian cvs Montecarlo (P. vulgaris) and Streamline770 (P. coccineus) were utilized for further studies. The joint axes between
cotyledons and embryos, a tissue rich of meristematic cells with high
aptitude to differentiate, were used. Abundant shoots were produced from
the explants in presence of different concentration of BA and/or TDZ. The
results showed that the absence or presence of cotyledons (a rich source of
nourishment and growth stimulation) is a determining factor to affect
individual tendencies and characteristics about the number of buds
(Mendoza de Gyves 1995).
A first attempt of regeneration from somatic tissues was made starting from
leaf-derived calli induced on a medium containing 2,4-D. After transferring
on liquid secondary medium with 2,4-D replaced by NAA, KIN, ABA and
GA3 the embryo-like structures reached the heart stage but did not became
plantlets (Allavena et al. 1994).
Focusing the efforts on P. coccineus, a species that gives fertile offspring
when crossed to P. vulgaris, direct somatic embryogenesis and
organogenesis mediated by small glossy calluses were obtained from
immature cotyledon explants on a modified half-strength MS medium
containing various concentrations of growth regulators.
The availability of an efficient protocol for plant regeneration from somatic
tissues and the susceptibility of the species or cultivar to infection by
Agrobacterium are required for successful transformation (Parson 2000). A
compatible reaction between Phaseolus and Agrobacterium was
demonstrated in our works. The presence of a detectable level of nptII
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activity in calli derived from several bean genotypes confirmed that
kanamycin resistance can be utilized as selectable marker in experiments
aimed to obtain transgenic plants (Allavena et al. 1994).
A home-made high-velocity microprojectiles bombardment device was
constructed and set up in Soressi’s lab. The efficiency of biolistic gene
delivery was tested in
cotyledons and embryo
axes of immature and
mature bean seeds using
plasmid pBI121, carrying
the ß-glucuronidase (GUS)
gene under the control of
the CaMV 35S promoter. In
(a) Transient GUS expression on Phaseolus
P. coccineus, 400±178
embryos after two days of bombardment
units transiently expressing
using tungsten microparticles covered with
the GUS gene were found
DNA. (b) Stable expression on Phaseolus
per shot. Recovery of
shoots after one month of infection with
transgenic
plants
was
Agrobacterium. In both images, arrows point
attempted by bombarding
to blue spots showing the expression of GUS
(Mendoza de Gyves 1995).
the apical meristem of both
P. coccineus and P.
vulgaris embryo axes to avoid cell re-differentiation from somatic tissues.
On average, 60% of the meristems showed at least one GUS expressing unit
after three consecutive bombardments on the same target (see Figure).
Chimaeric shoots expressing the GUS gene were sporadically found by
histochemical assay (Allavena et al. 1994; Mendoza de Gyves 1995).
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Bean breeding
Andrea Allavena (CRA - Research Unit for Floriculture and Ornamental Species,
Sanremo, Imperia Italy)
Bruno Campion (CRA - Research Unit for Vegetable Crops, Montanaso Lombardo,
Lodi Italy)
At the beginning of 70th Prof. Salamini together with Prof. Soressi set up a
robust breeding project mostly committed to develop bean cultivars
resistant to bean common mosaic virus (BCMV) and suitable for both direct
consumption and processing (canning and
freezing). Two were the procedures adopted:
crossing and selection; seed mutagenesis
induction. Along the first way with Fadda and
Allavena collaboration, the I gene conferring
hypersensitivity resistance to BCMV was
transferred from USA cv. Royal Red Kidney
(RRK) to Borlotto-type local varieties. Mary,
Fig.1 Seed-coat colour
Bea, Inge were the first BCMV resistant
mutations from RRK
varieties released in Italy (Salamini et al. 1978).
red seeds undergone to
With the second approach, the RRK red
EMS treatment.
coloured dry seeds were treated with EMS
mutagen and M2 populations screened for
seedling, plant and seed mutations (Fig. 1;
Motto et al. 1975). One of these white seedcoat mutants, after 1 backcross to RRK, led
to release BCMV resistant, Lisa P71 (Motto
et al. 1979b). By inter-crossing the best
BCMV resistant Borlotto bush breeding
lines and subsequent selection, new BCMV
resistant varieties improved for agronomic
and market characters were obtained:
Giulia, Lena and Minia (Fig. 2; Allavena et
al. 1982a). Then, a new series of BCMV
resistant varieties were developed: white
Fig.2 Borlotto-type bush
seeded Montebianco and Niveo, by interplant at waxy pod maturity
varietal crosses (Allavena et al. 1985a);
of the BCMV resistant cv.
Montalbano, marble white seeded and
“Lena”,
suitable
to
Mogano, beige seeded, by EMS treatment
mechanical harvesting.
(Allavena et al. 1985b). A new breeding
programme, started in 1975, was focused on
the introgression of the genetic tolerance to Pseudomonas phaseolicola
(from the PI150414 accession) into the best BCMV resistant lines issued
from the previous programmes. Highly tolerant offsprings were
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backcrossed to BCMV resistant lines and selected for resistance to both
diseases following artificial infection with both the pathogens. Monviso and
Morena were released as the best performing
bush breeding lines for fresh and dry grain
products (Allavena and Soressi 1982; Allavena
et al. 1982b).
Other halo blight partial resistant lines were
Montecarlo and Monterosa (Allavena et al.
1989). Grazia and Patrizia (Fig. 3) were new
bush, BCMV resistant, yellow-podded
cultivars, developed from [Meraviglia di
Venezia x (RRK and Kinghorn wax)] crosses’
breeding lines (Allavena et al. 1986). The
breeding work started by Prof. Salamini and
Fig.3 Extremely bushy
Soressi in 1971 for dwarfing P. coccineus
plant with yellow pods
of the BCMV resistant
through inter-specific cross with P. vulgaris
cv. Patrizia.
went
on
with
Campion, leading
to the development and release of the white
seeded Venere and the mottled seeded Alarico
cultivars (Figg. 4 and 5; Campion et al. 1981;
Campion 1995). Moreover, the Salamini and
Soressi research group studied seed size
Fig.4 Seeds of the cv.
inheritance in a
Venere.
cross of wild and
cultivated common bean (Motto et al. 1979b),
performed growth analysis in a reduced leaf
bean mutant (Motto et al. 1979a), contributed to
understand sucrose metabolism (Odoardi et al.
1976a) and detected multiple forms of
Fig.5 Seeds of the cv.
phosphoesoisomerase enzyme in bean seed
Alarico.
during its development in the pod (Odoardi et
al. 1976b). Carbohydrates metabolism was also investigated in smooth and
wrinkled pea developing seed (Odoardi et al. 1976c).
bitter fruit (bt)
see Glycoalkaloids
blind (bl)
see Monostem for tomato processing
Bowman Birk gene
see Helicoverpa armigera
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brown seed (bs)
see Tomato, genetic markers
Bruschetta
see Tomato, “fruit colour genes”
Cladosporium fulvum-2 (Cf-2)
see “Lesion mimic” mutants
Cystein proteinase inhibitor
see Helicoverpa armigera
DDRT-PCR
Giuseppe Testa, Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
Differential display RT-PCR has been adopted to study differencies in gene
expression in two experimental systems. In garlic, explants have been
compared before and after the induction of in
vitro proliferation in order to identify genes
involved in adventitious sprouting. The results
have been included in the PhD thesis of Dr.
Paula Bima (Bima 1997).
In tomato, RNA populations extracted from
ovaries of the parthenocarpic fruit (pat)
tomato mutant and from the corresponding
near isogenic wild-type were compared. The
sequencing and the characterization of three
Subepidermic placental
transcripts that were differentially displayed
expression of GAD3 in
in pat mutant ovaries compared to the wildthe tomato ovary (Testa
type
were described in detail (Testa et al.
et al. 2002).
2002). The protein predicted by the fulllength cDNA reconstructed from Clone 66 showed homology to peptides
encoded by self-incompatibility alleles of Papaver rhoeas and had a strict
ovary specific expression. The up-regulation seen at fruit set was an event
specific to the expression of parthenocarpy, because it was also found in
ovaries of a parthenocarpic fruit-2 line, a mutation which is not allelic to
pat.
The putative polypeptide deduced from the cDNA reconstructed from
Clone 72 showed high homology with plant H2A histones and preferential
transcription in ovaries.
A third fragment (Clone 91) was identical to the 3’ region of GAD3, a
tomato mRNA sharing similarities to short-chain alcohol dehydrogenases.
RT-PCR-based expression analysis revealed that this gene is preferentially
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transcribed in ovaries, sepals and leaves. The tenet that this enzyme family
has important functions in developmental pathways controlled by
hormones, and in particular gibberellins, is supported by its increased
transcription at the time of fruit set, by its positive response to treatments
with gibberellin A3, and by its expression localized in the mitotic cell layers
of the placenta (see Figure).
Dieffenbachia spp.
Enrico Santangelo (Università della Tuscia, Viterbo Italy)
Claudio Carrai (ARSIA, Pisa Italy)
Dieffenbachia is an ornamental species of relevant importance grown in
Europe and Usa. A five-years research program was set up in order to
define the heritable basis of leaf variegation and create new varieties by
means of intervarietal and interspecific crosses.
To make available a crossing
procedure efficient for genetic
investigations
and
breeding
programmes in Dieffenbachia spp.,
some experiments were carried out.
The degree of receptivity of the
stigmas and the fertilizing capacity of
the pollen were studied in relation
with different flower phenological
stages. Attention was also given to
seed extraction and conditions
favourable to seed germination and
seedling growth. The crosses
Dieffenbachia ripening fruits
involved the commercial cultivars
(Carrai et al. 1990).
‘Alix’, ‘Camilla’, ‘ Candida’,
‘Compacta’, ‘Jeanette’, ‘Katharina’, ‘Marianne’, ‘Nelly’, ‘Tropic white’,
‘Veerle’. On some of these cultivars both selfing and sib-cross were
accomplished. Fruit setting, seed germination and albino seedling
segregation were evaluated (Carrai et al. 1990; see Figure). Usually the
environmental conditions of the greenhouse where the Dieffenbachia is
grown hamper the flowering, that remain sporadic, poor and progressive.
Flowering has no value in the commercial production of plants, but it is of
basic importance for breeding programmes. With the purpose of increasing
the number of flowers and synchronizing their emission, the effect of GA3
treatments was studied (Fioravanti et al. 1991). Two different periods of
GA3 application (December and February) and four cultivars with different
ploidy levels (8n and 4n) were tested. All the cultivars responded positively
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to the treatment and showed an interaction between treatment period and
genotype for the number of flowers per plant and their concentration of
emission. The number of days from GA3 application to flower appearance
was lower in December than in February. The ploidy level of the genotype
seemed to influence both cross compatibility and response to GA3
treatments (Carrai et al. 1993). The number of flowers induced by GA3 was
dependent on plant age, vigour and branching.
easy peeling (ep)
see “Fruit skin” mutants
Ethylene
Enrico Santangelo, Valentina Fonzo, Mariagrazia Antonelli, Maurizio Enea
Picarella (Università della Tuscia, Viterbo Italy)
Although considered as the “ripening hormone”, ethylene plays a role in
many metabolic processes. The tomato “never ripe” group includes a
number of mutants with a direct or inderect impact on the synthesis and/or
perception of ethylene, displaying a phenotype with delayed and incomplete
ripening (Soressi 1975). Soressi’s activity has been concentrated on the
study of the tomato “never ripe” mutants Nr, rin and nor2, encompassing
also aspects of plant
physiology
other
than fruit ripening.
Studies of such
mutants have been
developed during the
70’s (see “Never
2
nor
rin
WT
ripe” mutants).
The availability of
Germination of 5-year-old seeds from nor2, rin and
WT 3 days after sowing.
such ethylene-related
genes in the same or
different backgrounds has extended concerns towards other physiological
aspects as seed germination and the role of ethylene in response to some
abiotic stresses. In Nr, rin and nor2, a more prompt and higher germination
percentage associated with a lower level of ethylene emission per seed
compared to the WT was observed (Lai et al. 2002, XLVI SIGA Annual
Congress; see Figure). By taking into account that 20 year-old seeds of
these ethylene mutants are still able to germinate, it can be speculated that
in these genotypes ethylene accumulation during seed senescence occurs
slowly and some processes, in which other hormones like auxin are
involved, are somehow altered during germination.
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Considering the role of ethylene as a signal molecule, the study of its effect
following treatments with abiotic constrains like high temperature,
luminosity, ozone and salt has been undertaken. The presence of Nr, rin and
nor2 genes has altered and modulated the leaf necrotic lesions occurring in a
tomato line displaying acropetalous autonecrosis at high temperature and
increasing luminosity (Fonzo et al. 2003, XLVI Annual SIGA Congress;
see “Lesion mimic” mutants).
When Nr and rin mutants have been exposed to a single, relatively high
dose of O3, an abiotic elicitor in plants, the results suggested that the
relationship between ethylene emission and the severity of O3-induced
foliar symptoms might be less straightforward than expected (Antonelli et
al. 2005, XV Meeting of the Eucarpia Tomato Working Group; see Ozone).
Finally, research on salt tolerance started in the first 90’s, has recently
involved the use of the nor mutant with the aim of analysing the interaction
between ethylene and abscisic acid (Picarella et al. 2007; see Salt
tolerance).
Fagiolo Badda
see Landraces, bean
Fagiolo del Purgatorio
see Landraces, bean
Fen gene
Mariagrazia Antonelli, Enrico Santangelo, Giorgio Mariano Balestra, Leonardo
Varvaro, Margherita De Biasi, Stefania Astolfi (Università della Tuscia, Viterbo
Italy)
In tomato, the resistance to race 0 of the phytopathogenic bacterium
Pseudomonas syringae pv tomato (Pst) is
governed by Pto, a member of a clustered gene
family that also includes the Fen gene that
confers sensitivity to the organophosphorous
insecticide Fenthion (see Figure). Therefore,
tomato cultivars that contain Pto also exhibit a
HR-like response to the insecticide. Other
components of the Pto resistance signaling
pathway are the Prf and PtiI genes.
In vitro regeneration on a medium containing
increasing doses of the insecticide was
Necroses on leaves of
employed to obtain regenerants resistant to
tomato cv. Rimone
Fenthion. Cotyledon explants of the hybrid
after Fenthion spray.
derived from the cross Rimone (resistant to Pst
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and sensitive to Fenthion) and Riogrande (resistant to Fenthion and
sensitive to Pst) were used. Santangelo et al. (1998) reported that the
methods allowed to induce variation in the Pto-Fen-Prf genic complex and
the selection of R2 plants with a different degree of tolerance to both the
agents (bacterium and/or insecticide) (Santangelo et al. 2000).
In order to explore the functional relationships among the expression of the
Pto/Fen gene family and the GOX transgene (see Glucose oxidase, GOX),
four tomato NILs (Riogrande and Rimone and their respective GOXtransformants, RC332 and RC131) were exposed to Fenthion, evaluating
the production of H2O2 and the activities of ROS-scavenging enzymes in
foliar tissue. Accumulation of H2O2 and transient enhancement of ROSrelated enzymatic activities could together confirm that Fenthion does elicit
an oxidative burst in the tissues of the tested tomato lines driven by either
Fen gene and GOX transgene. The Fen gene conferred sensitivity to
Fenthion, regardless of the expression of GOX. A prolonged accumulation
of H2O2 was observed in the leaves of Rimone and RC131, which was
instead transient in Riogrande and in RC332. In all the tomato lines,
exposure to Fenthion induced rapid transient changes in the activities of
most enzymes, except for peroxidase activity in the leaves of Rimone and
RC131, that steadily increased until the end of the sampling period. It is
suggested that the sensitivity of Rimone to Fenthion might be due to the
sustained activity of a H2O2-forming peroxidase (De Biasi et al. 2003). In a
study involving the tomato autonecrotic mutant V20368 (see “Lesion
mimic” mutants), treatments with Fenthion were performed to determine
whether the insecticide induced foliar lesions. Though the V20368 line
possesses the recessive pto and fen alleles, a necrotic phenotype mimiking
autonecrosis was observed after Fenthion treatment evidencing a lower
threshold of stress perception of this tomato line (Santangelo et al. 2003a).
The appearence of necroses on V20368 after Fenthion treatment was
confirmed by trypan blu staining followed by stereomicroscopic
observations (Antonelli et al. 2006, XIII SIPAV Annual Meeting).
“Fruit skin” mutants
Antonella Portesi (Università Cattolica Sacro Cuore, Piacenza Italy – presently
different address)
Piero Frangi (Fondazione Minoprio, Como Italy)
Giorgio Chioccia (Università della Tuscia, Viterbo Italy)
The tomato fruit, highly variable in weight and shape, according to
genotype and growth conditions, is endowed with a protective skin or
exocarp. Beneath this outer epidermal layer, there are 2 to 4 layers of
hypodermal cells with thicker walls and collenchyma-like thickenings. A
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layer of cutin is upon the thin (4-10 μm) cuticle, that covers the epidermal
cells. Cutinization extends into the radial walls of epidermal cells and may
also involve the hypodermal ones. The trichomes developed from epidermis
disappear at fruit maturity. Heavy cutinization contributes to the fruit
cracking resistance. Genotypic differences in pericarp and skin anatomy
exist and some of them are
of possible interest in
tomato
breeding.
The
mutants up to date described
are: sticky peel (pe), peach
a
b
c
d
(p), easy peeling (ep), waxy
Fig.1 Cuticole morphology of pe2 and p2
fruit (pe-2) and nonmutant fruits compared with cv. Gimar
ripening-3 (nor3).
WT: breaker (a) and red ripe (b) stages
pe gives crack-resistant
of the p2 mutant; red ripe stage of pe2
fruits but with a shiny,
mutant (c) and WT (d) (Portesi 1987).
extremely elastic and thin
skin, possibly lacking of
hypodermis. pe-2, mimicking pe, appeared in cv. Manapal. The p fruit has a
dull, opaque surface with increased hairiness when still immature. The ep
phenotype is caused by the disintegration of parenchyma cells with
intercellular air spaces below the hypodermis, thus permitting an ‘easy
peeling’ of the fruit. This mutant is of possible use in tomato breeding for
domestic or industrial peeled products. nor3 is a spontaneous mutation
appeared in the Italian “Tondo liscio di Pescara” landrace with colorless
and dull skin fruit, slow in ripening (Soressi 1975). Studies on the genetic
relationships among pe, pe-2, p and nor3 revealed that pe-2 is allelic to pe
and that nor3 is allelic to p. As a consequence and according to the TGC
rules the new symbols are pe2 instead of pe-2 and p2 instead of nor3 (Portesi
1987; Portesi et al. 1987).
Observations by scanning electron
microscope and histological analyses
have evidenced a clear-cut difference
in the cuticle of the mutants: it results
practically absent in pe2, thicker and
likely rugose (dull) in p2, thick and
smooth on the WT cv. Gimar (Fig. 1).
Such differences are recognizable from
Fig.2 Gimar fruit with p2
an early stage of fruit development.
phenotype, showing precocious
Likely, quali-quantitative variations in
joint-pedicel abscission.
cuticle waxes produced by the
epidermal fruit cells are involved. Another feature peculiar to p2 mutant is
an early abscission of the fruit at the pedicel joint (Fig. 2) in relation with
water availability. In addition the p2 fruit is resistant to cracking disorder
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on-vine and slowly shrivels during storage due to a higher permeability of
its skin to gases (CO2, O2, ethylene) and water. The p phenotype can also
ave some interests in tomato breeding by reducing fruit water content, with
lower pathogenic fungi attacks and also resulting in cracking resistance. To
avoid an early abscission of the fruit, it would be valuable to study the
combination of p2 with jointless mutants (j, j-2), thus improving the
performance against the mechanical harvesting shakes. The lower water
content of p mature fruits, besides requiring less heating power in
concentrating tomato puree, makes them particularly suitable for preparing
the Viterbese “Panzanella” food, made by spreading tomato red pulp onto a
slice of dry bread, previously soaked in fresh water, and pouring on it a spill
of extra-virgin olive oil.
Genetics and breeding
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
Despite his preference for following lab research and field trials, Soressi
was also engaged in writing contributions of general and/or didactic
interest, in the field of the teaching of genetics (Bianchi et al. 1973), plant
breeding (Soressi 1986a; Blanco and Soressi 1988), genetic resources
conservation (Maggioni and Soressi 1992) and agronomy (Soressi et al.
1981; Caporali and Soressi 1993).
Gibberellins
Andrea Mazzucato, Irene Olimpieri, Francesca Siligato (Università della Tuscia,
Viterbo Italy)
The role of gibberellins (GAs) in the development of male and female
tomato floral organs has been addressed in the study of the parthenocarpic
fruit (pat) phenotype (see parthenocarpic fruit). Whereas early research
showed that GA was able to rescue the anther phenotype but not
parthenocarpy in the mutant (Mapelli et al. 1979), a revision of the sensivity
of the mutation showed that gibberellic acid (GA3) was only effective in
restoring carpelloid anthers to the wild-type phenotype (Mazzucato et al.
1999).
Further reports indicated novel pat phenotypes that parallel those reported
in plants repeatedly treated with gibberellic acid or having a GAconstitutive response. The retained sensitivity to the GA-biosynthesis
inhibitor paclobutrazol revealed that this condition is dependent on GA
biosynthesis. Expression analysis of genes encoding key enzymes involved
in GA biosynthesis shows that in normal tomato ovaries the GA20ox1
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transcript is in low copy number before anthesis and only pollination and
fertilization increase its transcription levels and, thus, GA biosynthesis. In
the unpollinated ovaries of the pat mutant,
this mechanism is de-regulated and GA20ox1
is constitutively expressed, indicating that a
high GA concentration could play a part in
the parthenocarpic phenotype. The levels of
endogenous GAs measured in the floral
organs of the pat mutant support such a
hypothesis. Collectively the data indicated
Polycotiledony of a pat
that transcriptional regulation of GA20ox1
seedling, a phenotype
connected with altered
mediates pollination-induced fruit set in
GA synthesis and/or GA
tomato and that parthenocarpy in pat results
response regulation.
from the mis-regulation of this mechanism.
As genes involved in the control of GA
synthesis (LeT6, LeT12 and LeCUC2) and response (SPY) are also altered
in the pat ovary, it is suggested that the pat mutation affects a regulatory
gene located upstream of the control of fruit set exerted by GAs (Olimpieri
et al. 2007).
Glycoalkaloids
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
The tomato, as almost all species of the Solanaceae family, produces
glycoalkaloids (GLA, alpha-tomatine and dyhydrotomatine), secondary
plant metabolites that show toxicity against vertebrates. The desirability of
such metabolites in the tomato fruit is still a debated question; if GLAs are
of concern for consumers because of their general cellular toxicity to
vertebrates, they have proven to exert toxicity towards bacteria, fungi,
viruses and even arthropods, thereby serving as a potential source of
resistance to plant pathogens and pests. Recently, it was demonstrated that
assumption of alpha-tomatine in the mice diet is 20-fold less toxic than that
of other glyco-alkaloids and that the molecule also shows antitumoral and
anti cholesterol action.
Genetic knowledge on the control of GLA levels is fragmentary. In wild
cherry tomatoes, two accessions with bitter ripe fruits were found in some
Peruvian districts and the bitterness of their fruits was related to very high
content of GLAs (Rick et al. 1994).
In Italy, a variant with highly bitter fruit was discovered in a survey for
somaclonal variation in the background of cv. UC82 (Cunico, 1991; GP
Soressi, unpublished results). The trait, that was given the provisional name
of bitter fruit (bf), proved to be recessive and monogenically controlled
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(Cunico 1991). Although the causative relationship between the bitter taste
and a high tomatine content was not definitely proved, the phenotype
strongly recalls an impairment in GLA composition and represents a
valuable material being nearly isogenic to UC82.
Glucose oxidase, GOX
Riccardo Caccia, Stefania Astolfi (Università della Tuscia, Viterbo Italy)
Transgenic tomato (Solanum lycopersicum L.) plants containing the Gox
gene encoding glucose oxidase from Aspergillus niger were obtained in two
near isogenic lines (NILs) except for the Pto resistance gene (Riogrande
+/+ and Rimone Pto/Pto). T3 transgenic plants possessing a single Gox
gene copy were tested for resistance to Pseudomonas syringae pv. tomato
to ascertain the role of H2O2 in plant
pathogenesis response (Caccia et al.
1996, XL SIGA Annual Congress).
Reactive oxygen species such as H2O2 are
generated
following
pathogen
recognition, and they act as both cellular
signalling
molecules
and
direct
antimicrobial
agents.
The
Gox
transgene
Different blue color intensity
was regularly inherited and expressed in
of leaf disks from transgenic
the tomato plants. The level of GOX
Gox left) and control (right)
activity was one- to five-fold higher than
plants
following
an
in the non-transgenic WT (see Figure).
hystochemical assay.
The H2O2-generating, trans-Gox plants
proved to confer to the susceptible NIL the capacity to respond actively to
P. s. pv. tomato by reducing the number of viable bacterial cells in the
inoculated leaves and to limit the disease symptoms in terms of number and
size of leaf necrotic spots (Caccia 1989; Caccia et al. 1999; Balestra et al.
2000). To deeply explore the activation modes and timing of the plant
enzymatic systems involved in biotic and abiotic stress responses, the
effects of the Gox gene were studied by considering the interaction between
the Pto and Fen genes in the abovementioned NILs, taking into
consideration that the Rimone line is susceptible to the Fenthion molecule
based insecticide (see Fen gene). These transgenic NILs were also
investigated for assessing the damage level caused by polyphagous spider
mites (see Tetranychus urticae).
green flesh (gf)
see Tomato, “high-pigment genes”
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Helicoverpa armigera
Stefano Speranza, Riccardo Caccia, Valentina Fonzo, Elia Poerio, Claudio Pucci
(Università della Tuscia, Viterbo Italy)
Helicoverpa armigera (Hübner) is a cosmopolite Lepidoptera Noctuidae
able to injury several plant species among which tomato. The young larva is
yellow with black
head and, at the
mature stage, it
becomes
greenyellowish
with
typical lateral light
stripes.
Fig.1 Life cycle of Helicoverpa armigera (Hüb.).
It accomplishes its
life cycle in about a month,
but in Central-Southern Italy,
it performs 2-4 generations
per year, over wintering as
pupa in the soil (Fig. 1). In the
last years, this noctuid has
provoked infestations to
tomato of so high degree to
become a key pest in Central
Fig.2 Tomatoes damaged by H. armigera.
Italy and in several other
world regions (Speranza
2001). On the tomato plant,
the larvae start eating the
leaves, then feed themselves
with flower buds and berries,
by causing their dropping to
the ground or rotting (Fig. 2).
At harvest (August in Italy),
non
marketable
fruits
exhibiting holes and tunnels
without or with eating larvae,
Fig.3 Inhibition of cystein proteinase
can be observed.
activity in the raw extract of H. armigera
Proteinase inhibitors are able
larvae according with increasing quantity
to interfere in several animal
of proteins extracted by BG24 (—) and
digesting processes and to
RIG (—) leaves.
horn on their growing
(Brodway and Duffey 1986; Ryan 1990). Thus, plant proteinaceous
proteinase inhibitors have potential for increasing resistance of crop to
insect pests. With the aim of obtaining tomato transgenic plants resistant to
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37
H. armigera larvae, we transformed - the cultivar Riogrande with the gene
KTI3 (coding for a soybean Kunitz Inhibitor) and the cultivar UC-82 with
the gene Pi-IV (coding for a soybean Bowman-Birk Inhibitor). Leaf
extracts of transgenic plants contained significant levels of inhibitory
activities towards both bovine pancreatic trypsin and insect trypsin-like
enzyme, on average 187- and 5-fold higher-than controls, respectively. The
trypsin inhibiting activity was found stable at different plant phenological
phases in fruits and leaves; this activity resulted stable for 24 h in detached
leaf discs used to feed insects. Both inhibiting activities were able to
contrast development of H. armigera larvae, by interferring with midgut
trypsin-like activity that was found higher in III-IV ages than other ages
(Caccia et al. 1999, work presented at the workshop “Resistenza durevole a
stress biotici nelle piante: contributo delle biotecnologie”, Tuscia
University, Viterbo, 20-21 May 1999; Schettino 2004).
The introduction of the CpTI (cowpea trypsin inhibitor) gene into tobacco,
showed that its expression in the transgenic plants resulted in a significant
reduction of damage by another lepidopteran Noctuidea Heliothis virescens.
The mean weight was generally higher when larvae were fed with BG-106
transformed leaves. No statistical difference in mortality was observed
between larvae reared with control (RIG) and with Atcys transgenic tomato
(BG-106) leaves. The percentage of adults emerged from the cocoon was
81% and 76% for the control and BG-106 respectively. The sex ratio
(males/females) was in favour of the female sex both for the RIG (0.94) and
BG-106 (0.79) cocoons. On average, the number of laid eggs of the BG-106
fed females was 33% lower than the control. By considering the percentage
of hatched eggs (emerged larvae), the value obtained was 6.8% for BG-106
against 11% for RIG. According to these data, in Atcys transgenic tomato a
level of cystein proteinase inhibition double than the untransformed control
is sufficient to negatively influence the H. armigera biological cycle in
particular on number of eggs laid and on fertility (percentage of emerged
larvae per female), even if the weight of the larvae fed with the BG-106
leaves is on average higher than the control (RIG). The last datum is in
agreement with similar experiments reported in literature where the effect
of proteinase inhibitors is tested in different host-pest systems (Ryan
1990).
Tomato plants transgenic for genes coding for protein inhibitors, were
tested in order to evaluate their capability to inhibit the commercial
proteinase activities as well as those present in the midgut of the
phytophagous insect H. armigera. Leaf extracts of Pi-IV (gene from
soybean coding for a Bowman-Birk-like trypsin inhibitor) transgenic T3
plants (cv. UC-82) were able to inhibit bovine pancreatic trypsin and
trypsin-like activity of the phytophagous insect four times more than
control plant extracts. Extracts of KTI3 (gene from soybean coding for a
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Kunitz-like tripsine inhibitor) transgenic T3 plants (cv. Riogrande) were
able to inhibit bovine pancreatic trypsin and trypsin-like activity of the
insect 200 times more than control plant extracts; leaf extracts of AtCys
(gene from Arabidopsis coding for a cysteine protease inhibitor) transgenic
T2 plants (cv. Riogrande) were able to inhibit commercial papain and
cysteine-like activity of the insect four times more than control plant
extracts (Dirollo 1998). On the basis of the data it would be interesting, for
both speculative and applicative purposes, to use these transgenic plants in
order to verify in vivo the action on the mortality and/or on the reproduction
ability of the H. armigera, particularly dangerous to the tomato crop
(Farisei et al. 2002, XIX National Congress of Entomology, Catania 10-15
June 2002).
Aim of a further work was to perform a bioassay on the cotton bollworm,
feeding the larva with Kunitz proteinase inhibitor coded by the KTI3 gene
transgenic plant leaves. The molecules belonging to the Kunitz family have
a molecular weight of 21-24 kDa and inhibit the serine proteinase, resulting
active in the pH interval 9-11, that is the distinctive interval of Lepidoptera
larva midgut. Cv Riogrande tomato plants were transformed with the KTI3
gene through Agrobacterium tumefaciens. The analysis of the data obtained
and their elaboration show a direct negative effect on the Lepidoptera
biology, particularly in the second generation; this result could have, in the
future, a direct containment action of the phytophagous populations in
nature, causing a relevant decrease of the marketable production cull
percentage (Fig. 4; Speranza et al. 2002 XIX National Congress of
Entomology, Catania 10-15 June 2002).
high pigment (hp)
see Tomato, “high-pigment genes”
high pigment-2 (hp-2)
see Tomato, “high-pigment genes”
Induced variation, physical and chemical
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
Francesco Salamini (Parco Tecnologico Padano, Lodi Italy)
Soressi’s research activity begun in the early ‘60s, when he worked on his
thesis on the effect of X rays as mutagens for tomato seeds (Soressi 1964).
The first publications described this mutagenic approach considering
different genotypes and irradiation dosages on traits such as germination,
seedling height, time to flowering and ripening, plant height, fruit size and
number of seeds per fruit (Bianchi et al. 1963; 1964). The effect of gametic
and of intrasomatic (diplontic) selection was described in detail (Bianchi et
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39
al. 1963). The first mutants derived from these experiments were described
in detail, such as gamosepalous (fused sepals), typically sterile (aborted
inflorescence), skin finely wrinkled (rough fruit epidermis), cabbage leaf-2
(large and blistered leaves), finely netted (leaves paler and netted) (Soressi
et al. 1966; Cravedi 1966; Soressi and Cravedi 1967).
Soon after, experiments with chemical mutagenesis were started; treatments
with ethylmethanesulphonate (EMS) were used on San Marzano and cv
Sioux seeds and the general effects on the traits mentioned above described
(Marchesi and Soressi 1965). Interesting mutants were described such as
variegated (then renamed ghost-2, leaves with regions pale-green of
yellowish), turbinate corolla (petal tips turned), lazy-2 (stems prostrated),
reticulate virescent-2 (leaves yellowish with green veins), wrinkled leaflets,
male sterile-42 (potato leaf male sterile), short anthers (reduced anther
segments, see parthenocarpic fruit), hp-2 (high pigment) (Soressi et al.
1966; Cravedi 1966; Soressi and Cravedi 1967; Cravedi and Soressi 1969;
Bianchi and Soressi 1969; Soressi 1970a; Soressi 1975).
Later, the effects of combining chemical mutagenesis with in vitro induced
variation was studies (Mensurati 1994, see Induced variation, somaclonal)
In Phaseolus vulgaris, dormant seeds were treated with different doses of
EMS and several variants in chlorophyll content and seed morphology and
colour obtained. The M1 plants were chimerical, often having only a small
mutated sector. Based on the segregation ratios obtained, the number of
initial cells present in the apical meristem ranges from three to eight for the
whole plant and from one to three for the primary branches. The
experimental results indicate a post-treatment origin of the primary branch
initials. No evidence was found for the occurrence of diplontic selection
during the ontogeny of a mutated plant in the variety studied (Motto et al.
1975).
Induced variation, somaclonal
Francesco Mensurati (Università degli Studi della Tuscia, Viterbo Italy)
Antonella Portesi, Cristina Cunico, Elena Perri, Giuseppe Stancanelli (Università
Cattolica del Sacro Cuore di Piacenza, Italy)
Roberto Ranieri (Stuard experimental farm, Parma Italy)
The genetic changes occurring in somatic cells during adventitious
meristem formation and shoot regeneration is referred as somaclonal
variation. The first approach for setting up the regeneration procedure, with
Soressi’s supervision, started in the 80’s at the Piacenza Botany and Plant
Genetic Institute (Antonelli et al. 1987; Portesi et al. 1989, XXXIII SIGA
Annual Congress, Sassari). Perri (1991) studied the effect of culture
medium composition, explant type and the starting genotype on the
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frequency of regenerants and induced variations. Cunico (1991) described
and characterized the most interesting somaclones in particular the first
“bitter fruit” (bf) mutant, singled out in S. lycopersicum. A contribution for
innovating nursery in horticulture is reported in Soressi et al. (1991). To
have an effective comparison between the segregant progenies of the
regenerated plant and its true control plant, the rescue of the cotyledon
explant donor seedling was done by in vitro rooting of the root-deprived
hypocothyl.
Following the cv. Alice cotyledon explants regeneration, at the University
of Parma, the percentage of R1 progenies segregating mutants, including
seedling lethals, was 11% of the control and 48% of the R2 from
regenerated somaclones (Tiburtini 1992). The unusual frequency of twins
(from polyembryonic seeds) observed in the R2 progenies, was attributed to
an hormonal influence during the sexual embryogenesis in the R1
regenerated plant (Cunico et al. 1992, XXXVI SIGA Annual Congress).
A further experiment (1988-1989) in collaboration with the University of
Parma and the support of the Stuard farm was accomplished by plant
regeneration from hypocothyl cotyledon and leaf callus of cvs. Alice and S.
Marzano. The data confirmed somaclonal variation as a powerful tool for
creating genetic variability (Ranieri et al. 1996). By excluding polyploids,
26% of Alice progenies and 9% of S. Marzano showed mutations which
were consistent through generations. Among the chlorophyll mutants the
mottled green fruit shoulder (um) was described for the first time.
Certainly, highly informative data on somaclonal variation came from the
PhD thesis of F. Mensurati (1994). In this work, by recovering the
cotyledon explant donor seedlings, the mutagenic effect of EMS seed
treatment and cotyledon regeneration of EMS treated and non treated seeds,
was evaluated in a monostem (to-2) line and in the cv. Alice. The procedure
adopted permitted to compare, per each genotype, the distinct segregating
progenies from the control (R0S1), the EMS treated seeds (M2), the
cotyledon regenerated control (R2) and the regenerated cotyledons of
seedlings from EMS treated seeds (R2M2). The main results are dealing
with the M1 and R1 plant chimaerism, the low (5%) frequency of polyploid
phenotypes among the regenerated plants, the mono-Mendelian recessive
mutant frequency that resulted significantly lower in the cotyledon explant
regeneration than that following EMS seed treatment. The combination of
EMS seed treatment with regeneration of cotyledon-derived seedlings from
EMS-treated seeds allowed to recover mutants that would otherwise be lost
by merely selfing the M1 plants. This combined procedure succeeded in the
screening of a cv. Alice anthocyaninless mutant induced by EMS seed
treatment, but recovered through its cotyledon regeneration. In fact, in the
literature on tomato somaclonal variation via regeneration no
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41
anthocyaninless mutant has been found in opposition to that occurred
following EMS seed treatments (see Figure).
Specimen of recessive mutants recovered in segregating progenies
following cotyledon regeneration, EMS seed treatment and their
combination: sublethal yellow and WT seedlings, a; sublethal twins from
polyembryonic seeds, b; adult plant of the bitter fruit (bf) mutant with
yellow calyx trait, c; seedlings without (left) or with (middle and right)
anthocyanin, d.
In vitro culture in vegetable species
Enrico Santangelo, Riccardo Caccia, Emilio Mendoza-de Gyves (Università della
Tuscia, Viterbo Italy)
Paula Bima (Facultad de Ciencias Agropecuarias, Universidad de Córdoba,
Argentina)
In the present Laboratory of Biotechnology of Vegetable Species at
DABAC, tissue culture has become a routine approach for breeding
Fig.1 Tomato shoot regeneration from
hypocotyl explants.
Fig.2 Tertiary axillary shoot
emerged
from
nodal
microcutting;
arrows:
cutting sections of the
primary and secondary
shoots previously formed.
purposes. The contributions here cited
played a relevant role in achieving not
negligible results. Efforts were made to develop more efficient
methodologies by refining culture media for plant micropropagation,
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42
regeneration and transformation of different plant species such as tomato,
garlic, bean, chickpea and others. Moreover, in vitro cultures were utilized
also in basic research.
Most of the micropropagation, cotyledon explant regeneration and
transformation procedures applied, also involving physiological analyses,
were carried out on tomato.
Sources of tolerance to abiotic stresses were screened among shoots of
genotypes differing in salt sensitivity regenerated on media with various
NaCl concentrations (Habashy et al. 1993, XXXVII SIGA Annual
Congress). Plants tolerant to Pseudomonas syringae pv. tomato not
showing the typical necrotic leaf spots due to Fenthion spray, were induced
by regenerating shoots on media containing this insecticide molecule
(Santangelo et al. 1995-ISPAV, Roma, Italy).
Chemical mutagenesis through EMS seed treatment coupled with cotyledon
explant regeneration and rescue of the apical meristem from the cotyledonexcised seedlings were successfully employed for increasing the range of
somaclonal variants obtainable in tomato (Mensurati et al. 1995, XXIX
SIGA Annual Congress). GA3 supplied to culture medium was effective in
inducing axillary meristems in shootless monostem tomato (to-2) (Bima et
al. 1995). To reduce the polyploids frequency among the shoots regenerated
in media supplied with hormones, the capacity of hypocotyl explants to
regenerate in hormoneless media was assessed, confirming that both
regenerating ability and polyploids frequency are highly dependent on the
genotype, not omitting that hypocotyl is a mixoploid tissue (Fig. 1; Bima et
al. 1993). On the basis of such indications, to develop triploid tomato
plants, the tetraploid parents crossed with the diploid ones have been
obtained not by colchicine treatment (Allavena et al. 1976) but through
hypocotyl explant regeneration (Habashy et al. 2004).
Cotyledonary explant regeneration and vegetative propagation via nodal
microcuttings were steps routinely adopted in our experiments of genetic
transformation (Schettino et al. 1998).
In tomato, each in vitro nodal microcutting produces only one axillary
shoot; aiming to obtain supernumerary axillary shoots per nodal
microcutting, different media, with or without plant growth regulators, were
tested. Both IAA and zeatin were able to gradually induce the proliferation
of supernumerary axillary shoots when the previously formed were
removed. To avoid the occurrence of polyploid variants among the plants
derived from supernumerary shoots, auxin (IAA) is to be excluded in all the
steps of shoot proliferation. The sole cytokinin (zeatin) when added to the
basal medium gave the highest proliferation efficiency (Fig. 2; Cammareri
2003; Soressi et al. 2008).
Efforts for in vitro organogenesis and embryogenesis in Phaseolus vulgaris
were made by Allavena and Rossetti (1983, 1986). In the 90’ s in many labs
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numerous experiments were planned in bean to set up tissue culture and
genetic transformation protocols. To investigate their morphogenetic
potentiality, apical meristems, from 24 h water-soaked dry seeds, were
induced to form adventitious shoots in presence of 1-2 mM of thidiazuron
(TDZ). The data obtained confirm a competence of P. coccineus better than
P. vulgaris even dependent on the intraspecific variability; the best
proliferating genotypes tested resulted to be cv. Venere and cv. Montecarlo
respectively (Ranalli et al. 2001).
The need for a reliable in vitro vegetative propagation protocol in garlic
encouraged P. Bima, tutored by Soressi, to set up an efficient methodology
for micropropagating garlic clones of different origin (Bima 1997).
Insect resistance
Stefano Speranza (Università della Tuscia, Viterbo Italy)
The interest of GP Soressi towards insect resistance begun in the early ‘90s
when experiments aimed at exploiting the introgression of morphological
characters from wild species into cultivated potato were carried out
(Remotti 1990). Later, this topic was expecially developed in tomato where
the genetic transformation with protease inhibitor genes (Kunits, Bowman
Birk and cystatine) was tested as a method for conferring insect resistance.
A long and fruitful collaboration with Prof C. Pucci and Dr S. Speranza of
the Department of Plant Protection at the Tuscia University allow to test the
effect of this genetic improvement towards the tolerance against important
tomato noxious insects (see Helicoverpa armigera; see Spodoptera
littoralis).
Kunitz gene
see Helicoverpa armigera
Landraces, bean
Lucia Lioi (Istituto di Genetica Vegetale, CNR, Bari Italy)
Bruno Campion (Unità di Ricerca per l’Orticoltura, CRA, Montanaso Lombardo
Italy)
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
From the hillside to the shores surrounding the Bolsena lake (Viterbo),
within the Municipalities of Gradoli and Acquapendente, since long ago has
been cultivated the bean landrace “Fagiolo del Purgatorio”, so called
because traditionally dined during the Purgatory luncheon of Ash
Wednesday. This crop covers an area of ∼ 20 ha, yielding 8-10 q/ha of dry
grain. The research activity, carried out in the framework of the MIURetic
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44
MIPAF project “Valorisation of the Italian vegetable germplasm” and of a
research contract with the Comunità Montana Alta Tuscia Laziale, aims to
characterize both the bean plant and its product together with the
phytosanitary healing. Bio-agronomic trials were developed by comparing
the seed accessions provided by local farmers. On the initial seed samples
and on those reproduced in open field at Viterbo, biochemical (seed storage
proteins), nutritional and molecular-genetic (SSR and AFLP markers)
analyses were carried out (Santangelo et al. 2006; Lioi et al. 2007). The
results revealed the extent of genetic diversity in this bean landrace and
permitted to select homogeneous, representative accessions. Moreover they
have evidenced the Meso-american gene-pool origin of the “Fagiolo del
Purgatorio” and the presence of two plant growth habits (bush determinate,
bush indeterminate) highly correlated with the seed storage protein
phytohaemagglutinin (PHA) electrophoretic variants SG2 and MG2
respectively. The two distinct growth habits differ in plant height, number
of nodes and pods, pod length, seeds per pod, earliness and dry grain yield
and in addition in sensitivity to pathogens.
In collaboration with the “Gian Pietro Ballatore” Consortium and in the
framework of the “Piano per la produzione di proteine vegetali (3PSicilia)”, a morphological and molecular analysis has been carried out on
the landrace from Sicily “Fagiolo Badda”. Compared with other Italian
landraces, this characteristic bi-coloured bean showed a very late flowering
time and a ripening of pods in October. Molecular analyses (ISSRs)
indicated that the Badda bean belongs to the Andean gene-pool and is
distinguishable from other borlotto types or bi-coloured varieties (Paniconi
2006).
Landraces, lentil
Andrea Mazzucato, Pietro Mosconi (Università della Tuscia, Viterbo Italy)
Research on lentil has been recently carried out in collaboration with two
public consortia. The Comunità Montana Alta Tuscia Laziale committed a
molecular characterization of the landrace “Lenticchia di Onano” from
Northern Latium, whereas the “Gian Pietro Ballatore” Consortium in the
framework of the “Piano per la produzione di proteine vegetali (3P-Sicilia)”
commited the characterization of three landraces from Sicily (“Lenticchia
di Linosa”, “Lenticchia di Ustica”, “Lenticchia di Villalba”). Several
accessions were fingerprinted by ISSR markers; all the landraces clustered
separately (Gianfilippi 2006). For the “Lenticchia di Onano” landrace, the
molecular analyses confirmed the suspicion that actually two different
morphological and molecular typologies are cultivated and commercialized
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as “Onano” landrace. Only further molecular and historical investigation
could reveal which is the true original type.
Landraces, tomato
Andrea Mazzucato, Giorgio Chioccia (Università della Tuscia, Viterbo Italy)
Research has been carried out on several tomato landraces with particular
focus on those cultivated in Latium such as Scatolone di Bolsena, Pantano
romanesco, Spagnoletta del golfo di Gaeta e Formia (Mazzucato et al.
2006).
Scatolone is a landrace cultivated on the hills surrounding the Bolsena lake
with very good organoleptic qualities and a characterisc boxiness
(scatolatura) of the fruit. An extended germplasm collection has been done
in 1994 and a mild selection has been done among 42 accession from ten
different farms (Pepponi 1995). The landrace is characterized by the
occurrence of vegetative, “male” individuals that correspond to the “rogue”
phenotype described in some British and Dutch cultivars; such character
shows a weak genetics basis and the only way to reduce the occurrence of
rogues is to keep low the germination temperature (Mazzucato et al.
1998a).
A selection of Italian landraces showing high differentiation for fruit shape,
size and utilization has recently been used to identify marker-trait
relationships in an association mapping approach (Mazzucato et al. 2008a;
see Association genetics). Also very recent is a collaboration with the CRAUnità di Ricerca per l’Orticoltura of Monsampolo del Tronto (AP) for the
molecular characterization of the “A pera abruzzese” landrace.
The Soressi’s group has been referenced by the Regional Agency for
Agricultural Development for the inclusion of tomato landraces from
Latium in the “Registro Volontario Regionale delle risorse genetiche
autoctone della regione Lazio”.
“Lesion mimic” mutants
Enrico Santangelo, Mariagrazia Antonelli (Università della Tuscia, Viterbo Italy)
“Lesion-mimic” mutants, mimicking cell death, are a powerful tool for the
comprehension of mechanisms activated during plant stress response.
Soressi singled out in a progeny under selection of Dr. Svetlana, Moscow, a
tomato plant (V20368) that, at increasing temperature and high light
intensity, exhibited leaf necrotic lesions, with acropetal progression. The
autonecrosis resulted due to the interaction of the Cf-2 resistance gene
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(from Solanum pimpinellifolium) with Rcr3esc gene, encoding for a cysteine
protease, of S. lycopersicum (Santangelo et al. 2003a).
The mutant revealed to be a valuable tool for studying molecular processes
common to different stressing agents. The Cf2/Rcr3esc interaction is already
active in asymptomatic leaves of
the autonecrotic line V20368,
causing in turn a strong oxidative
burst (Santangelo et al. 2003a;
Santangelo et al. 2007a). A delay,
or a lacking, in the activation of
the defensive barrier has been
observed by Marabottini et al.
(2007,
LI
SIGA
Annual
Congress):
scavenging
enzymes
of
Sympomatic leaves of the V20368
H
O
(catalase)
were
functioning,
2
2
autonecrotic line (Santangelo et al.
but not all those committed to
2003a).
superoxide removal; the system of
ascorbate-glutathione cycle was at levels of non-stressed controls; markers
of PCD and HR were both activated during the autonecrotic process.
Chloroplasts and mitochondria appeared directly affected by oxidative
stress.
Considering the pivotal function of ethylene signalling in response to
stresses, Fonzo et al. (2003, XLVI SIGA Annual Congress) investigated its
role in modulating the necrosis formation by crossing the V20368 line with
the ethylene tomato mutants Nr, rin and nor (see Ethylene). An ethylene
burst resulted evident when the autonecrotic process was triggered in
V20368, but in the F1 hybrid with the WT plant modifications in some steps
of the ethylene metabolism altered and modulated the phenotypic
expression of the autonecrosis. A consistent maternal effect appeared, thus
supporting the hypothesis of an involvement of mitochondria in the
triggering and spreading of the autonecrosis.
In V20368, Santangelo et al. (2007b) observed a remarkably higher level of
HCN associated with an ethylene burst and supposed an overwhelming of
the detoxifying capacity of β-CAS (enzyme of mitochondrial fraction
detoxifying cytosolic CN-) and AOX1b (mitochondrial alternative oxidase)
that led to an inhibition of mithocondria respiration, ROS overproduction
and cell death.
When treated with ozone, the mutant line V20368 (but not its control)
developed, 96 h after the fumigation, O3-dependent necrotic lesions
associated with a significant increase in both foliar ethylene release and
CN- levels (Antonelli et al. 2007). Finally, the histochemical analysis of
V20368 leaves treated with Fenthion revealed, the presence of dead cells
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(absent in the control RioGrande) thus confirming its greater sensitivity to
abiotic stresses (Antonelli et al. 2006, XIII SIPAV Annual Meeting).
lutescent-2 (l-2)
see Tomato, “high-pigment genes”
MADS-box genes
see parthenocarpic fruit; stamenless-2
Maize, hybrids
Carlo Lorenzoni (Università Cattolica del Sacro Cuore, Piacenza Italy)
Compared to that in tomato or other vegetables, Soressi’s activity in maize
genetics and breeding has been limited. In a report dated 1972, comparison
data for 725 opaque-2 hybrids are presented. The yield trials were located
in the Po Valley at Cremona, Castelnuovo B.A. (Milano), Pozzuolo
(Udine). As a result of these experiments, the authors identified some
pedigrees that reduce the well known disadvantages of the opaque-2 maize
(Maggiore et al. 1972).
Male sterility
see 7B-1 mutant; pistillate; positional sterile-2; stamenless-2,
parthenocarpic fruit
Microgravity
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
In a colaborative research with the group of Prof. F. Saccardo, a horizontal
uniaxial clinostat has been used to grow tomato plants under simulated
microgravity conditions. The study was carried out to evaluate the
vegetative and reproductive behaviour of the dwarf tomato variety ‘MicroTom’ in simulated microgravity conditions and to determine if tomato
plants would complete their life cycle (‘seed-to-seed’). Plants grown under
simulated microgravity exhibited a spreading growth and an increased
internode length. Total fruit yield, leaf area, leaf dry weight, fruit dry
weight, total dry weight and shoot/root ratio were lower in the clinorotated
tomato plants than those grown in the control treatment. The amount of
foliar carotenoids, and chlorophyll a and b were also substantially reduced
under simulated microgravity conditions. Quality parameters (total soluble
solids and fruit dry matter) of tomato plants were also negatively affected
by clinorotation. Although the number of flowers was increased, fruit
setting was reduced by 46% under clinorotation. However, no significant
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difference was recorded for the pollen fertility and the seed number per
fruit. Seeds produced by clinorotated plants were germinated proving to be
biologically and functionally normal, thus indicating that ‘Micro-Tom’
plants could realize complete ontogenesis, from seed to seed, in
microgravity (Colla et al. 2007).
Monostem for tomato processing
Bruno Campion, Massimo Schiavi (CRA- Research Unit for Vegetable Crops,
Montanaso Lombardo, Lodi Italy)
Piero Frangi (Fondazione Minoprio, Como Italy)
Giuseppe Gandolfi (Università Cattolica del Sacro Cuore di Piacenza, Italy)
According to Soressi, the persistent problem of processing tomato
mechanical harvesting in Italy during the the ‘60-‘80s could be got over by
putting in practice the monostem
ideotype represented by a short,
possibly erect plant without
lateral shoots, bearing only one
truss with 3-5 firm fruits, to be
grown at high density population
(Soressi 1974, 1975a, 1976).
Such an ideal plant should result
in a tomato crop with an early
“simultaneous”
agronomic
Fig.1 High density field plot with
ripening
due
to
the
short
monostem plants at fruit turning stage.
flowering period and the
reduced number of fruits per plant counterbalanced by a high number of
plants per unit area. By trying to select such a kind of plant, the known
mutants lacking or having few lateral shoots were collected and
characterized. Some of them, namely
blind (bl), lateral suppressor (ls),
torosa-2 (to-2), eramosa (era),
unicaulis (uni), were crossed with
some breeding lines and commercial
cultivars (Petomech, CaJ, XP1030).
Up-right plants, reduced in size,
without lateral shoots, bearing 1 truss
with 1-2 seeded fruits, therefore close
to the monostem ideotype, were
Fig.2 Monostem early, ripe plant of
found in the F2, from the crosses
the 2nd selection cycle.
involving only bl or to-2. Plants with
analogous habit werw found in the F2 progeny segregating ls but because of
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their smaller fruits, variable in seed number and size, they were not
considered. Following crosses between the best to-2 lines and in few cases
backcross to seven commercial cultivars, different monostem (STU) lines
were chosen. On the basis of an allelismo test (Schiavi et al. 1980), to-2 and
bl mutants resulted to be attributable to same locus and their expression
influenced by the background effect. The F2 to-2 plants judged particularly
interesting for the programmed breeding purposes were crossed and
backcrossed to 13 commercial processing varieties. Among these prebreeding stocks, the first 17 S4 monostem (STU) lines were selected
(Schiavi, 1977) and agronomically evaluated for earliness, concentrated
ripening, adaptability to high density population (Fig. 1) in comparison
with the cvs. Ventura and UC105 (Campion 1977). Some of them were
used for studying growth parameters (LAI, LAR, NAR, HI, SL, DW) in
comparison with cv. UC82 (Frangi 1984; Frangi and Soressi 1986) and
ones as NILs for investigating physiological and biochemical aspects
(Soressi and Mapelli 1992). The same monostem line was used to evaluate
its response to various plant population density, ranging from 2.75 to 8.8
plants/sqm (Soressi et al. 1986). The breeding activity went on by selecting
the progenies from the inter-crosses of the best monostem lines (Fig. 2)
derived by the previous crosses with 13 commercial varieties (Bruzzone et
al. 1984). The build-up and distribution of the dry matter and some of its
components in both monostem and normal genotypes were also examined
(Chiappari 1993). Potential and limits of the monostem type to be exploited
for the tomato breeding was analyzed by Scolari (1987) who evaluated
earliness and quality of the raw and processed product. In order to increase
the number of flowers/fruits per truss, the interaction of to-2 and s
(compound inflorescence) genes was examined in segregating progenies,
remarking an epistatic effect of to-2, so impairing next efforts. Altogether,
three cycles of selection were carried out by combining the best monostem
lines from the 2nd cycle. At the end of the 3rd selection cycle some of the
best performing STU lines were evaluated during 1988-90. Gandolfi (1992)
characterized the most interesting monostem (STU) breeding lines, giving
also their pedigree through 4 cycles of subsequent selection. All these
breeding lines are earlier (10-15 days) than Red Setter, with equal or
superior quality (pH, ° Brix and Bostwick) but inferior for yield. Therefore,
none of the breeding lines selected was suitable for wide field cultivation in
relation to the high plant density needed (250-300 thousand/ha). For
reducing such a high density plant population, an higher fruit number per
plant (from 2-3 to 4-5) should be required. However, some indications have
emerged about the possibility of using the earliest monostem lines for a 2nd
tomato field culture (after pea or barley) without yield reduction. A parallel
selection work followed crosses between to-2 and pat-2 (Poma 1992).
Poma contributed to select, evaluate and describe a number of monostem
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(to-2), parthenocarpic (pat-2) lines of possible interest for both table and
early processing tomato.
Monostem in protected cultivation
Pardossi Alberto, Tognoni Franco (University of Pisa, Italy)
The breeding work with
monostem breeding lines for
open field cultivation prompted
Soressi to elaborate a new plant
model for energy saving in
greenhouse
(Soressi
and
Scarascia Mugnozza 1988) and
for growing tomato plants
particularly suitable to soilless
culture techniques (Soressi et al.
Monostem, non parthenocarpic, poor
fertile
line
(a),
corresponding
1988). Taking into account the
parthenocarpic pat-2 line with improved
difficulties of pollen dehiscence
fertility and fruit size (b).
and fertilization in flowers of
monostem plants in NFT culture (Pardossi et al. 1988), in collaboration
between Tuscia University and Pisa University, glasshouse trials with
monostem parthenocarpic (pat-2) lines were carried out by adopting the
NFT culture system. In fact, in Mediterranean winter-spring condition, the
pat-2 gene interacts favourably with to-2 not only improving flower set but
also increasing fruit size (Baroni 1994). Positive features, constraints and
perspectives are reported in Soressi et al. 1992.
“Never ripe” mutants
Enrico Santangelo, Maurizio Enea Picarella (Università della Tuscia, Viterbo Italy)
The tomato “Never ripe” group includes the mutants Never ripe (Nr),
ripening inhibitor (rin),
non-ripening (nor) and
nor2 (Soressi 1975b)
with an altered synthesis
and/or perception of
ethylene, displaying a
Fruits of wild type (+/+) and ripening mutants at
phenotype with delayed
full ripening.
ripening process and a
modified fruit pigment synthesis and production. The dominant mutation
Nr encodes for an ethylene receptor, determining the production of fruits
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that do not fully ripen. rin results from a deletion in the last exon of a
tomato MADS-box transcription factor gene designated LeMADS-RIN; the
ripening process is blocked before the climateric burst and the fruit has a
slow ripening becoming lastly light yellow. The nor locus harbors a gene
with structural features suggestive of a transcription factor. The nor2
mutant, allelic to nor, had been collected in 1968 by Soressi in a vegetable
garden of a grower who considered it as a “falling-in-love” type for an
unexplicable (to him) occurrence of a plant with red fruits among the
yellow-orange nor2 “winter type”. Both rin and nor genes behave as
incompletely recessive for both fruit colour and biochemical level. Indeed,
while positively affecting the fruit shelf-life, these heterozygous in
opposition to the homozygous genotypes, do not affect so negatively the
fruit colour. In Italy, the first biochemical investigation on fruits of the
“never ripe” mutants (Nr, rin and nor2) and high pigment (hp and hp-2) and
green flesh (gf) has been promoted by Soressi (Martiniello 1975; Palmieri
et al. 1975; Palmieri and Soressi 1978). In these papers the authors stressed
the theoretical and practical interest in particular of hp-2 and nor2 to raise
the vitaminic content and to naturally extend fruit shelf-life and storage
without loss of nutritional value. The influence of Nr and mainly nor2
alleles on quality and storage of fruits from different table and processing
cultivars and their F1 hybrids with the same two mutants was studied by
Soressi’s ISPORT group and IVTPA, Milan (Palmieri et al. 1976; Rossi
1977; Sozzi et al. 1981). A key contribution in this topic is represented by
the two field experiments in F. Rossi’s dissertation (1976) with fruit storage
trials at room and controlled (12°C, 80-90% R.H.) conditions. Productive
and quality field data revealed the nor2 allele affecting, even when
heterozygous in F1 hybrids, the shelf-life of the on-vine tomato fruits, so
allowing a greater ripening concentration of commercial yield in once-over
harvest. Shelf-life and storage values, colour, lycopene, beta-carotene,
chlorophyll, sucrose, reducing sugars and vitamin C contents of 9 F1
hybrids (4 Nr/+ and 5 nor2/+) were reported in Palmieri et al. (1976) and
Sozzi et al. (1981). These authors put in evidence notable advantages
expected by exploiting the nor2 heterozygote in F1 tomato hybrids suitable
to mechanical harvesting and processing with high quality fruits. The effect
of the nor2 mutant in Gimar background on the fruit quality during on-vine
ripening, over-ripening and post-harvest was investigated in Lancellotti’s
graduation thesis, then published in Allavena et al. (1978). The recorded
traits were: days from flowering to green mature, turning stage, ripening
and four subsequent overripening stages; °Brix, dry matter, acidity, and the
same abovementioned traits (total carotenoids, etc.). These authors stated
that the better shelf-life, firmness and quality of the Gimar nor2/+ fruits,
either on-vine or post-harvest, offers encouraging perspectives for delaying
in time and space the consumption of fresh fruits or their processing.
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On the basis of such results and perspectives discussed with Soressi, Nira
Retig supplied with nor2 seeds, stimulated breeders from Israel to develop
F1 tomato hybrids, heterozygous for nor2 or rin, producing firm, longstoring fruits, both on-vine and post-harvest. Naomi (Hazera Seed
company) was the first F1 nor2/+ cherry tomato hybrid, rapidly spread and
grown from Sicily to the North of Italy. These Israelian nor2/+ and rin/+ F1
hybrids changed the way of harvesting tomato, from picking fruit by fruit at
turning stage, into truss by truss with red fruits, thus opening the era of the
long shelf-life red-ripe tomatoes.
non ripening (nor)
see “Never ripe” mutants
non-ripening3 (nor3)
see “Fruit skin” mutants
Onion
Massimo Schiavi, Bruno Campion (CRA- Research Unit for Vegetable Crops,
Montanaso Lombardo, Lodi Italy)
At the beginning of the 1970’s the authorities of Parma Province supported
the formulation of a breeding program for the onion variety “Dorata di
Parma” by the Institute of Botany and Plant Genetics, run by Prof.
Lorenzoni, at Piacenza University. The aims were to solve the main
problems found in the populations grown locally (high sensitivity to
Fusarium basal-rot and lack of bulb uniformity). The Emilia Romagna
Region’s Department of Agriculture also recognized the importance of this
crop in its territory and agreed a separate program for onion genetic
improvement with the Experimental Institute for Vegetable Crops of
Montanaso Lombardo (LO) under the Soressi guide. Considering the
similarity of the two programmes, both the Institutes recognized the
benefits from joining the two forces and requesting the collaboration of the
Institute of Plant Pathology, University of Bologna, that had already begun
studies on onion phytopathological constraints. Afterwards, the Italian
Ministry of Agriculture increased the funds for the onion programme within
the “Orticoltura” project (Soressi and Porcelli 1988), sub-project
“Asparagus and onion” (Soressi 1988a; 1988b).
The selection methods adopted were: mass selection on naturally infected
soil and three systems of recurrent selection with evaluation of S1, S2 and
half-sib progenies both on infected soil and by means of artificial infection
(Schiavi et al. 1984d; 1984e; 1988a). Within these activities a different
pronubous efficacy of honey bee towards blowfly in controlled onion
pollination was ascertained (Fig. 1; Schiavi et al. 1988b). After three cycles
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53
of selection the improved populations showed a percentage of rotten bulbs
twice lower than that of the commercial
populations of Dorata di Parma and
similar to tolerant commercial hybrids
(Fantino and Schiavi, 1987). Progress
was also observed in bulb characteristics:
uniformity of shape, colour, shelf life and
scale covering. In addition, some
researches were carried out to study
methods
for
onion
androgenesis
Fig.1 Caged blowflies for
(Campion et al. 1985, XXVIII SIGA
onion selfing.
Annual Congress), here reminding
Campion’s later success with
onion gynogenesis. At the end of
the breeding program, five new
cultivars were released: “ERSO
1”, “ERSO 2”, “Marica” and
“Morgana” (Fig. 2) of the type
“Dorata di Parma” and “Sorriso”
derived from the “Borettana”
variety (Schiavi et al. 1991). Two
Fig.2 Morgana: an improved onion
graduation theses were produced
variety tolerant to Fusarium basal-rot.
on evaluation of different
techniques of onion seed production including “seed to seed” procedure:
Moretti (1982) and Donati (1980).
Ozone
Mariagrazia Antonelli, Enrico Santangelo (Università della Tuscia, Viterbo Italy)
Maurizio Badiani (Università Mediterranea di Reggio Calabria Italy)
Ozone (O3) is the most widely spread atmospheric pollutant in the
industrialized world. When O3
concentration
exceeds
the
tolerance threshold for a given
plant, it results in the formation
of visible damage, which has
been described as necrotic
lesions,
bifacial
flecking,
-1
One hundred nL L ozone for 5 h causes
bronzing, and "waterlogging".
visible symptoms in the lesion-mimic
At full development, these
mutant V20368 (left) but not in its
symptoms physically resemble
phenotypic control line MP-22 (right).
HR lesions. O3 requires entry
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into the apoplast through the stomata to exert its toxicity. Once inside the
leaf, O3 reacts with the cell wall, the plasma membrane and apoplastic
fluids, to produce a burst of reactive oxygen species, such as superoxide
radicals, hydrogen peroxide, and hydroxyl radical. Therefore, O3 behaves as
an abiotic elicitor in plants, thus suggesting the possibility of using such
photochemical oxidant as an experimental tool for studying plant defence
responses towards biotic and abiotic stresses. A fruitful collaboration
among the universities of Reggio Calabria, Pisa and Perugia, supported by
MIUR from 2003 to 2007, elucidated some molecular events and the
complex cross-talk among hormonal signalling routes, deployed in tomato
upon O3 exposure. Tomato mutant lines impaired in ethylene perception Nr
(Never ripe) or biosynthesis rin (ripening inhibitor) (see Ethylene), the
tomato autonecrotic line V20368 (see “Lesion mimic” mutants) and
anthocyanin mutants (see Anthocyanins) were used as model systems. In a
first contribution, it was showed that both the Nr and rin mutations affect
activation timing and relative transcripts abundance of the ethylene
biosynthetic enzymes 1-aminocyclopropane-1-carboxylic acid (ACC)
synthase and ACC oxidase, as well as of the ethylene receptor NR, in
response to realistically elevated O3. Since both mutants and their near
isogenic WTs were equally damaged by the O3 treatment, a sort of isoform
"compensation" among ethylene biosynthetic genes could be supposed,
with the common aim of producing an ethylene burst triggering defence
responses. Such a burst was indeed observed, and showed similar time
patterns in the three genotypes, although the peak of ethylene emission
levels were different (Antonelli et al. in press). Also the autonecrotic line
V20368 revealed to be a valuable tool for studying common molecular
responses to different stressing agents in tomato. In response to O3
fumigation, V20368 plants showed a high level of ethylene release and
increasing CN- level compared with the control line. Such results support
once more the link between an O3-dependent ET burst and the development
of foliar symptoms, and suggest that CN- overproduction might take part in
the launch of O3–induced PCD (Antonelli et al. 2006).
The anthocyanin atv mutant (see Anthocyanins) seems to be more resistant
to O3 than other tomato genotypes in terms of foliar symptoms, confirming
a role of anthocyanins as ROS scavengers.
‘Panzanella’
see “Fruit skin” mutants
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parthenocarpic fruit (pat)
Andrea Mazzucato, Luigi Selleri (Università della Tuscia, Viterbo Italy)
Dario Beraldi (University of Edinburgh, Edinburgh UK)
The parthenocarpic fruit (pat) mutant is the first source of parthenocarpy
(fruit set in absence of pollination) described in tomato and was obtained by
Bianchi and Soressi (1969) with mutagenesis
with ethyl methanesulfonate (EMS). The
mutant phenotype, characterised by a high
tendency to parthenocarpy and abnormalities
in anther development, was firstly ascribed to
the action of two tightly linked genes, sha
(short anthers) and pat (parthenocarpic fruit)
(Soressi and Salamini 1975). Data
demonstrating the practical interest of the sha
pat trait for selecting parthenocarpic varieties
to be cultivated in open field for processing
Fig.1 A fruitful triploid
are reported by De Logu (1974). The
parthenocarpic pat plant.
discovery of a spontaneous sha-pat mutation
in the line ‘Montfavet 191' (Pecaut and Philouze 1978) proved that the
described phenotype was caused by a single recessive mutation with
pleiotropic effects and the gene was finally named pat (Soressi 1985;
Philouze and Pecaut 1986). The
mutant shows an enhanced
ovary growth rate during the
first 10 days after anthesis,
which
correlates
with
a
precocious onset of cell
divisions in the pericarp and a
higher auxin, gibberellin and
DNA content in ovaries
Fig.2 Variety Partheno plant, showing
(Mapelli et al. 1978). At
the exuberant amout of ripe cherry-type
maturity,parthenocarpic
pat
fruits, mostly seedless.
fruits are typically about twothirds of the normal size (Falavigna et al. 1978), probably due to decreased
cell enlargement, rather than to a lower number of cell layers (Mapelli et al.
1978). Compared to the near isogenic WT, the mutant shows a general
better earliness and higher fruit set (Falavigna and Soressi 1987), and a
higher content in total soluble solids (Falavigna et al. 1978), therefore
raising practical agronomic interest.
In addition to parthenocarpy, the pat gene causes aberrations that affect
male floral organs, i.e. short, irregular and apparently unfused anthers that
leave the stigma exerted (Bianchi and Soressi 1969; Philouze and Pecaut
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1986). Moreover, while other sources of genetic parthenocarpy are usually
facultative (they also can produce normal seeded fruits; Philouze, 1985),
female fertility is greatly reduced in pat and seed production is more
difficult. More recent research showed that the female sterility of the pat
mutant is due to the aberrant development that affects a fraction of the
ovules (Mazzucato et al. 1998b) and to overall defects in pollen guidance
that occur in the ovary (Mazzucato et al. 2003).
Physiological aspects of the mutant phenotype have addressed the role of
plant growth regulators, such as polyamines (Antognoni et al. 2002; see
Polyamines) and gibberellins (Mapelli et al. 1979; Mazzucato et al. 1999);
studies on gibberellin regulation have been extended to a more general
perspective on fruit set (Olimpieri 2006; Olimpieri et al. 2007; see
Gibberellins). In addition, the pat system has served for studying the
interaction with the lateral suppressor mutation (Ziliotti 1984), for the
identification of transcripts differentially expressed in parthenocarpic
ovaries (testa 1998; Testa et al. 2002; see DDRT-PCR), to synthesize
polyploid parthenocarpic lines (Fig.1) (Allavena and Soressi 1978; Habashy
et al. 2004; see Polyploidy) and to study the relationship between
parthenocarpy and the activity of class B MADS-box genes (Mazzucato et
al. 2008b, see stamenless-2).
Parallel breeding work led to the constitution and inscription in 1982 of the
new variety “Partheno”, a cherry type homozygous for the pat gene, in the
National register of varieties (Fig.2).
pat is also the first mutation for parthenocarpy in tomato that has been
mapped; it is located on the distal end of the long arm of tomato
chromosome 3 (Mazzucato et al. 1998c; Beraldi 2002; Beraldi et al. 2004).
The ongoing positional cloning project has presently reduced the genomic
window containing the Pat locus to a couple of BAC clones, spanning
about 170 kb (Selleri et al. 2007, contribution to the LI Annual SIGA
Congress).
While the list of loci of the ‘pat series’ has nowadays reached the
description of pat-9 (Gourguet et al. 2008), the founder member of this
family still deserves much attention and promises interesting results to
researchers.
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parthenocarpic fruit-2 (pat-2)
Enrico Santangelo (Università della Tuscia, Viterbo Italy)
The fruit setting in tomato early production under cold plastic-tunnel is
strongly hindered in Mediterranean areas by low-night and wide fluctuating
temperatures, high light intensity and growing daylenght, environmental
conditions occurring during spring season. The introduction of
parhenocarpy genes into cultivars
of value may encompass such
constraints and avoid the use of
chemicals. One of the most
important source of parthenocarpy
is the parthenocarpic fruit (pat-2)
gene found in the Russian cultivar
Severianin and frequently used in
breeding programmes (Santangelo
and Soressi, 1990). The interest
for such a gene is due to its simple
heredity and its capacity to form
seedless fruits of comparable size
Mature fruits of a tomato hybrid in
respect the normal ones when the
his normal (+/+) and parthenocarpic
environmental conditions could
(pat-2/pat-2) versions.
hamper it, allowing a high stability
and uniformity of production. This mutation has been deeply studied in
France by the group of J. Philouze and in Spain by the group of F. Nuez.
In Italy, experiments were planned to study pat-2 expression and its effects
on productivity in conditions of temperature fluctuating during growth.
Different French hybrids were compared in their normal and parthenocarpic
pat-2 near isogenic versions (Santangelo 1988; Cantoni 1991). The pat-2
gene allowed an improvement of the fruit setting, when relatively low-night
temperature (1st truss) or high-day temperature (4th truss) occurred.
Therefore, more stable production and greater commercial fruit size
uniformity were obtained. These advantages were more or less pronounced
considering both microclimatic conditions or the genetic background where
the pat-2 gene is introduced (Bricchi 1993; Mapelli et al. 1995).
Because the average number of seeds per fruit never achieves that of the
correspondent normal version, such a traits represents a good parameter for
selecting parthenocarpic plants (Santangelo 1988; Santangelo et al. 1990).
An interaction between the hormonal systems governed by the two different
genic situations (WT pat-2) has been observed for the average number of
seeds per fruit and the mean fruit weight and diameter. When the
environmental conditions are favourable to pat-2 expression,
parthenocarpic seedless fruits of optimal size are produced. When the
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conditions are more advantageous for seed production, also the
parthenocarpic genotypes produce seed, but in a lower number, leading to a
light decreasing of fruit diameter.
Efforts for mapping the Pat-2 locus have been undertaken by using
segregating populations in purely cultivated genetic background (Marras
2002), after crosses with introgression lines (Olimpieri 2006) and with S.
pennellii plants (unpublished results).
Pathogenic microorganism culture filtrates for the induction of
aspecific plant resistance
Francesco Mensurati (Università degli Studi della Tuscia, Viterbo Italy)
Paola Crinò (ENEA, C.R. Casaccia, Roma Italy)
Tomato crop cultivations are still requiring chemicals applications to
control Phytophtora infestans, causing heavy loss of production, being
partial the genetic resistance available for this pathogen. In vitro selection
with toxins or culture filtrates of plant pathogenic microorganisms has
allowed, in some cases, to improve the efficiency of plant breeding
programmes by contributing to isolate disease resistant plants. The use of
toxins should be preferred to that of culture filtrates because they are pure
substances which do not cause artefacts and do not falsify the results of in
vitro selection (Crinò et al. 1996).
Under the Soressi’s guidance, and tutored by P. Crinò, Mensurati (1991)
attempted to increase through in vitro plant regeneration in presence of the
pathogen culture filtrate, the resistance level of tomato cultivars possessing
the Ph-2 gene which confers partial resistance to P. infestans.
The inoculation essays on seedlings or foliar discs did not evidence any
kind of resistance due to the Ph2 gene in comparison to the susceptible
genotype, lacking of such a gene. For the experiments planned with culture
filtrate or toxin for improving the plant response to P. infestans, the cv.
Piline was chosen for its relatively higher number of shoots per explant
regenerated in vitro. Therefore, 1290 cotiledonary explants of Piline were
cultured on selective medium containing the toxic metabolites of P.
infestans. After two subsequent transferrings on substrate containing fresh
culture filtrate, 10 non-susceptible plantlets of cv. Piline were in vitro
regenerated, rooted and propagated. The work proved a relatively fair
efficiency of the in vitro selection procedure in presence of toxic
compounds as a simple and rapid way for the identification of resistant or
tolerant plants.
peach (p)
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see “Fruit skin” mutants
Phaselieu Concerted Action
Enrico Santangelo (Università della Tuscia, Viterbo Italy)
Ali Akbar Habashy (Agricultural Biotechnology Research Institute, Karaj Iran)
The overall aim of the Concerted Action PHASELIEU was to coordinate
the on-going research on Phaseolus and to elaborate an integrated strategy
model for the improvement of Phaseolus production in Europe for human
consumption. The 13 partners participating to the project were
representative of Universities and Research Centers of Spain, France,
Germany, Belgium, Austria, United Kingdom, Portugal, Israel and Italy.
Prof Soressi was the responsible of the Italian group and spokesperson for
Italian “Phaseolus workers”: B. Campion (Istituto Sperimentale per
l’Orticoltura, Montanaso Lombardo Lodi), D. Bollini (CNR Biosynthesis
Lab, Milano) and P. Ranalli (Istituto Sperimentale per le Colture Industriali,
Bologna). Collaborative links with R. Papa (Università Politecnica delle
Marche, Ancona) and within the framework of the European Cooperative
Programme for Crop Genetic Resources Networks (coordinated by L.
Maggioni of Bioversity) were established. Within this collaboration, 403
accessions of the Phaseolus collection originally maintained at the
University of Cambridge were evaluated and regenerated, by coordinating
Italian institutions engaged in bean breeding or research.
Inside the Phaselieu network, the Soressi’s team focused its efforts on
setting up in vitro protocols for multiple shoots induction and genetic
transformation via Agrobacterium or particle gun (see Bean,
biotechnological approach). The morphogenic capacity of apical meristems
excised from dry seeds was evaluated by inducing them to form
adventitious shoots on media containing several combination of TDZ and
2,4-D (in P. coccineus, cv. Venere) or BAP (in P. vulgaris, cv.
Montecarlo). The results confirmed a higher regeneration competence of P.
coccineus than P. vulgaris. During 5-6 subcultures, 10-15 shoots per
explant were obtained for cv. Venere, whereas the cv. Montecarlo did not
exceed 4 shoots. The possible use of thin layers of cotyledonary nodes of P.
coccineus (cv. Venere) was also investigated. The results showed a good
capacity of proliferation allowing to obtain 8-9 shoots per explant after 4
subcultures and indicated this procedure as particularly suitable for genetic
transformation by Agrobacterium tumefaciens (Parsons 2000). Provided the
induction of multiple shoots from apical meristems in both P. vulgaris and
P. coccineus, the group investigated the potentialities of the biolistic system
for bean genetic transformation. In 11 independent experiments, 1178
explants were bombarded, whose 193 (16.4%) rooted in selective medium
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Multiple shoots induction in apical
meristems of cv SL770 (left) and
Venere (right) (Parsons 2000).
containing kanamicin and 15 (7.7%)
resulted positive to PCR (Santangelo
et al. 2000; V Phaselieu Workshop,
Hannover).
Soressi’s group contributed to the
five workshops held during the
Concerted Action and to the Final
Dissemination Meeting. The Third
Workshop on “Biotic and abiotic
stresses in bean” was held at Viterbo
on 4-5 June 1999.
pistillate (pi)
Andrea Mazzucato, Irene Olimpieri (Università della Tuscia, Viterbo Italy)
In this research, we characterized the pistillate (pi) mutant, a genotype that
directly recalls mutations affecting class B MADS-box genes, but that has
not been further studied after the first description in the middle of the last
century. Plants homozygous for the pi allele appear with Mendelian
proportions in segregating populations and, compared to wild-type, show
delayed flowering, a frequently modified sympodial segment, higher
occurrence of compound inflorescences, and reversion of floral meristem to
vegetative identity. The most striking
aberration in pi mutant flowers is the
homeotic transformation of stamens
into carpels. More rarely, homeotic
conversions are reported also in the
second floral whorl, with staminoid and
carpelloid
petals.
Ultrastructural
analysis reveals more or less subtle
sepaloid features in the three inner
Parthenocarpic fruit set on a pi
floral whorls, mainly based on the
plant showing a secondary
presence, distribution and amount of
ectopic fruit and sepal or shootglandular and non glandular trichomes.
like outgrowths.
In the ovary, a ‘flower within flower’
phenotype was seldom observed; in one instance such phenotype was
coupled with the setting of a parthenocarpic fruit, that reiterated the
differentiation of a new flower (see Figure; Olimpieri and Mazzucato,
submitted). Mapping information positioned Pi on the distal end of the long
arm of chromosome 3. According to published maps, this position is not
compatible with any of the known class B and E MADS-box tomato genes
(Mazzucato et al. 2008b; http://www.sgn.cornell.edu); thus the identity of
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Pi has to be searched among those genes that act upstream to the ABC(DE)
system of determination of the identity of floral organs.
Poliploidy
Andrea Mazzucato, Enrico Santangelo, Maurizio Enea Picarella (Università della
Tuscia, Viterbo, Italy)
Andrea Allavena (Res Unit for Floriculture and Ornamental Species, Sanremo Italy)
Ali Akbar Habashy (Agricultural Biotechnology Research Institute, Karaj Iran)
Polyploid crops are a valuable resource for horticulturists owing to their
peculiar advantages with respect to the diploid ones. In tomato, the
exploitation of either natural or artificially induced polyploidy turned out to
be scarcely promising for the high sterility of triploids and the low yield
and high variability in size of the autotetraploid fruits, extremely variable in
seed number which is positively correlated to fruit size in this species.
Soressi’s research on tomato polyploidy began in the 70’s when he first
obtained data evidencing the potential of polyploidy coupled with
parthenocarpy for tomato breeding. As reported elsewhere (see
parthenocarpic fruit and parthenocarpic fruit-2), parthenocarpy represents
a useful trait for genetic improvement
since it allows good yields in
environments
unfavourable
for
pollination and fertilization. Soressi and
co-workers (Allavena 1975; Allavena et
al. 1976; Allavena and Soressi 1978)
Representative
fruits
of
obtained tetraploid seeds from handdiploid (left), triploid (centre)
pollination of polyploid shoots following
and tetraploid parthenocarpic
colchicine treatment of WT and pat/pat
plants (Habashy et al. 2004).
F3 seedlings. These seeds produced 4x
plants either parthenocarpic, as fruitful as the corresponding diploid pat/pat,
or WT almost completely sterile in greenhouse. The 4x parthenocarpic
fruits were larger and heavier (30%) than their corresponding 2x pat/pat. In
addition, their soluble solids content and pH did not significantly differ
from the pat/pat diploids’, while the ascorbic acid content was higher
(20%).
Since hormonal treatments of spontaneous tomato triploids gave high fruit
yield and quality (Kagan-Zur et al. 1991), a programme to develop
genetically parthenocarpic triploid tomato plants through hand-crossing was
planned. To this aim Soressi’s team regenerated in vitro, from hypocotyl
explants, diploid and tetraploid somaclones belonging to lines carrying pat
or pat-2 mutantions and to the respective WTs. The yield recorded in open
field plots proved the parthenocarpic polyploid (4x) fruits to be larger than
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the parthenocarpic diploids (2x), despite their almost complete seedlessness
(Kiferle 1998; Habashy et al. 2000, V Giornate Scientifiche SOI). These
new 4x and 2x somaclones, in near-isogenic background except than for the
pat mutation, were hand crossed each other to obtain triploid (3x) plants. In
greenhouse, 3x pat/pat/pat plants yielded seedless and high quality fruits
differently from the non-parthenocarpic triploids, completely unfruitful and
sterile (Santangelo et al. 2003b; Habashy et al. 2004).
This unique, successful result offers a valuable contribution to practically
exploit polyploidy in tomato by combining it with genetic parthenocarpy.
To tomato breeders, it offers the challenge of the synthesis of fruitful
triploid F1 commercial hybrids, with high-quality, seedless, uniformly sized
fruits diversified between fresh consumption and processing. Such triploid
hybrids, by considering their complete apireny, shall necessarily be
vegetatively propagated. Furthermore, the triploid tomato genotypes, even
transgenic, unable to produce fertile pollen and seeds, will nullify the risk
of gene flow to conventional tomato cultivars (Santangelo et al. 2005, 2nd
Solanaceae Genome Workshop, Ischia, Italy).
Polyamines
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
Polyamine (PA) pattern was investigated in tomato floral organs plants at
four developmental stages from floral bud until 2 days after anthesis.
Differences in putrescine, spermidine and spermine content, both free and
conjugated, were evaluated in a wild-type line (WT) and in the
parthenocarpic fruit (pat) mutant. PA content was quite similar in WT and
pat mutant ovaries, but the two lines showed a different PA pattern
throughout flower development. A net accumulation of PAs was observed
in concomitance with pre-anthesis floral stages in pat mutant ovaries, while
in WT plants the PA pattern did not change significantly during the
developmental stages considered. These differences seem strictly related to
the physiological events that initiate ovary development, which have been
shown to be precocious in the mutant ovaries and occurring at pre-anthesis
stages (Antognoni et al. 2002).
A decrease in free PA levels and an increase in the conjugated pool were
associated with the initiation of cell expansion. The pattern of ornithine
decarboxylase, arginine decarboxylase and S-adenosylmethionine
decarboxylase gene expression, as well as the respective enzyme activities,
were also analysed during development. The lack of correlation between
transcript levels and enzyme activity suggests that post-transcriptional
control mechanisms must be acting in the regulation of biosynthetic genes
(Antognoni et al. 2002).
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Pollen-pistil interaction
see parthenocarpic fruit
positional sterile-2 (ps-2)
Fabrizio Ruiu (Università della Tuscia, Viterbo Italy)
For the production of F1 hybrids, the hybridization of tomato plants is done
by manual emasculation and pollination. This is a labour-intensive process
and does not guarantee 100% hybridity. There are many genic male sterility
systems available in tomato (Badino and Soressi 1976); however they are
generally not used on commercial scale because of difficulties with the
incorporation into seed parent lines and their maintenance. The recessive
mutation positional sterile-2 (ps-2) has been studied from this perspective
(Ruiu 2006). The flowers of male sterile plants do not differ from those of
isogenic male fertile
line except for the
dehiscence phenotype;
in the mutant, the
stomium does not open
properly and the pollen,
although viable, is not
released (see Figure).
Dehiscent anthers in the WT line WT (left) and
The mutant is female
functional sterile phenotype in ps-2 (right).
fertile.
The expressivity of male sterility can vary with the genetic background and
with the environmental conditions, but shaking of the inflorescences does
not increase seed yield (Ruiu 2006). This mutation offers the possibility to
maintain the female line through manual-selfing, enabling to obtain a
complete male sterile progeny. Furthermore, comparative studies conducted
in Bulgaria, where the mutation has been described for the first time, on the
time necessary for the emasculation of floral buds (as practiced when using
fertile seed parent) and flowers at anthesis (as practiced when using a ps-2
line as seed parent), made it clear that emasculation at anthesis was easier
and almost two times quicker than emasculation applied on floral buds. The
Ps-2 locus has recently been finely mapped by Gourguet et al. (2006).
potato leaf (c)
see Tomato, genetic markers
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Programmed cell death
Riccardo Caccia, Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
We investigated the senescence of flag leaves of durum wheat (Triticum
durum) during grain-filling, and of petal-like ray flowers of Jerusalem
artichoke (Helianthus tuberosus) at anthesis. In both systems, we observed
cleavage of DNA to high molecular weight fragments, followed by further
degradation to nucleosomal fragments (laddering), a classical hallmark of
apoptosis. We show that DNA fragmentation in such specialised leaves is
triggered
early
in
organ
development,
before
the
appearance of visual symptoms of
senescence. Our observations
support the hypothesis that
senescence and programmed cell
death (PCD) are part of the plant
Programmed cell death occurring in
developmental program, activated
the style transmitting tissue after
by developmental cues (Caccia et
pollination (left), compared with an
al. 2001).
unpollinated style (middle) and a
Further research addressed the role
DNAse-treated
positive
control
of PCD in reproductive tissues and
(right). Black spots represent nuclei
organs; TUNEL analysis on such
with PCD-controlled DNA cleavage.
tissues revealed a common
occurrence of PCD in the anther tapetum, in the nucellus and in the
transmitting tissue of the style after pollination (see Figure; unpublished
results).
Pto
see Glucose oxidase, GOX; Fen gene
Reporter genes
Marianna Rossi, Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
The characterization of mutants, whose phenotype is putatively determined
by altered hormone levels or perception may shed light on the role played
by hormones in different aspects of plant developmental biology,
physiology, biochemistry and molecular biology. In this research, the auxin
accumulation dynamic has been analysed in a number of tomato floral
mutations by means of a reporter construct where the auxin-inducible
promoter of the Agrobacterium tumefaciens gene 5 was cloned upstream of
the uidA gene (GUS) (Rossi 2002; Mazzucato et al. 2006). The mutants
selected for the analyses were: uniflora, lateral suppressor, stamenless 2,
fasciated, parthenocarpic fruit-2. All the mutants where crossed with plants
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harbouring the reporter construct in homozygosis and F2 families were
screened for the presence of the transgene (km resistance). From WT and
mutant plants segregating in the F2 families, plants organs were collected
and subjected to the histochemical GUS assay. Non transgenic specimens,
used as negative control, were always devoid of reporter signal. In turn,
remarkable, organ-specific differences were reported between mutant and
WT plant tissues.
WT roots showed strong auxin accumulation at the root tip and in
correspondence of the adventitious root primordia. The uf, sl-2 and f mutant
showed no difference from the WT at the root level. Differently, pat-2 and
ls roots showed a lower IAA content. Signal intensity was also overall
associated to root vigour. In leaf tissues, for the WT and all mutants the
signal was absent
or very weak, with
the exception of sl2 that showed
higher
IAA
content.
The WT ovary
IAA accumulation as detected by the p5::GUS
showed
an
reporter construct in the tomato root apex (left), the
secondary root emergence zone (middle) and the
increasing auxin
leaflet (right). The upper row is the negative control.
content through its
development into
fruit, up to a maximum that was reached at the mature green stage. While
the uf ovary did not show any difference from the WT, pat-2 and f ovaries
and their young fruits had a stronger reporter signal, in agreement with the
putative role of the hormone in parthenocarpic development and in carpel
number specification. However, auxin content in the pat-2 ovary was lower
than in the WT at later stages of fruit development, as it was in the partially
sterile ls mutants; such observation strengthens the knowledge that fruit
IAA is essentially contributed by the seeds. The reported results on floral
mutants and on other developmental tomato mutants have been the object
of a PhD thesis (Rossi, 2003).
The same reporter construct has been used to study the IAA-GA
relationship in plants silenced for the GA biosynthesis gene GA20ox-1
(Olimpieri and Mazzucato unpublished).
ripening inhibitor (rin)
see “Never ripe” mutants
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Salt tolerance
Stefania Astolfi, Maurizio Enea Picarella, Ali Akbar Habashy, Enrico
Santangelo, Sabrina Zuchi, (Università della Tuscia, Viterbo Italy)
Alberto Pardossi (Università di Pisa, Italy)
In the frame of Prof. Soressi’s work on plant response to biotic and abiotic
stresses, some experiments were dedicated to the salinity stress in tomato.
This crop, widely cultivated in the Mediterranean Region requires frequent
water supply during spring-summer and this may represent a drawback for
its cultivation especially in coastal areas (primary salinity) and greenhouse
(secondary salinity).
To
succeed
in
breeding
for
a
complex trait like
salt tolerance, the
availability of as
much reliable as
easy-to-score
parameters is crucial.
In vitro cotyledon
regeneration
was
coupled
with
salt
18S-rRNA-normalized, hybridization signal values,
stress to induce
obtained through northern blot analysis, of TAS14
genetic variability to
and TSW12 gene probes, on mRNAs derived from
seedlings grown in 150 mM NaCl. Values are
be screened for
relative to those from mRNAs of non-treated
sources of tolerance
seedlings.
to
this
abiotic
constraint. Nuclear
and cytoplasmic genome alterations were induced in seedlings and plants of
R2 generation. Attention was also paid to in vitro vegetative growth
parameters such as fresh and dry biomass of root and shoot of seedlings
grown on media with different NaCl concentrations (0, 50, 100, 150, 200
mM). Among the genotypes tested, cvs. Cal J and Edkawi proved to be the
most tolerant while cv. UC82 the most sensitive to salinity (Habashy 1992;
Habashy et al. 1993, XXXVII SIGA Annual Congress, Perugia, Italy). The
response to NaCl of these genotypes was confirmed in a further trial in
which cvs. Edkawi and Cal J performed better, respectively for root and
shoot biomass, than cvs. UC82 and Petomech. In parallel, a positive
correlation between such growth response and the expression of specific
salt-responsive genes (see Figure), was put in evidence (Picarella et al.
1995, ESF Network-Workshop, Maratea, Italy). Analogous results were
reached by an indipendent experiment on the same genotypes but
considering early growth rate of 21-days-old plants, grown in greenhouse
1200
Hybridization signal (pixel)
1000
800
TAS14 gene
600
TSW12 gene
400
200
0
Edkawi
etic
UC82
Petomech
CalJ
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with soilless system culture, as a tolerance-related parameter (Pardossi et
al., unpublished data).
Salt tolerance was also investigated in terms of physiological traits. In
particular, ethylene (ET), regarded as a key hormone regulating gene
expression of environmentally challenged plants, released from 4-week-old
plants grown in a versatile controlled hydroponic system, was measured
along with parameters connected with plant salt-stress response like
root/shoot biomass, chlorophyll content and others related to the plant
nitrogen metabolism, like inorganic osmolites (Cl-, NO3-), free aminoacids
contents and nitrate reductase (NR) activity (Astolfi et al. 2004, XXII SICA
Annual Congress, Perugia, Italy). Edkawi, as a salt tolerant cultivar, and cv.
Gimar (WT) and its two near-isogenic lines (NILs) carrying the nor2
(Gimar NOR) and Nr (Gimar Nr) mutations, respectively involved in the
synthesis and perception of ET (see “Never ripe” mutants) were compared
in control (10 mM NaCl) and salinity (70 mM NaCl) conditions. An
interesting result was the antagonistic relationship between NO3- and Cl-,
suggesting leaf nitrate content as a marker of plant NaCl tolerance. In fact,
the salt treatment caused a considerable increase of leaf NO3- content only
in the tolerant cv. Edkawi, but not in the Gimar NILs. Salinity, then, caused
a significant inhibition of NR activity in leaves of Gimar Nr, Gimar NOR
and Edkawi plants. In particular, the NR activity increased (+23%) in
Gimar WT and was reduced (-75%) in Edkawi, where the highest NO3accumulation was measured. Therefore, a very small amount of NO3appears sufficient for NR induction while this activity is not induced or
even inhibited when NO3- is beyond a certain threshold. Leaf chlorophyll
decline is generally exerted by salt exposure of plants, probably correlated
to an indirect effect of NaCl on the content of essential nutrients. In salttreated plants, whilst a significant decrease in leaf chlorophyll was recorded
for cv. Edkawi, Gimar WT and Gimar Nr NILs, a 33% increase was seen in
Gimar NOR (Astolfi et al. 2005, XXIII SICA Annual Congress, Turin Italy;
Astolfi et al. 2005). Unpublished data proved that the nor2 mutation does
affect not only the fruit ripening by suppressing the respiratory climateric,
but also the vegetative organs, by reducing ET release in leaf and root of
seedlings (4th leaf stage). Moreover, in salinity conditions, the nor2 mutation
has a beneficial influence on the content of antioxidants by increasing
ascorbate and tocopherol in Gimar NOR fruits, thus improving their
nutritional value (Sgherri et al. 2007).
As recently advanced by some authors, an increase of the abscisic acid
(ABA), another stress-responsive hormone, is required to prevent excess
ET production from plant tissue under drought stress. In such conditions
ABA synthesis would allow the plant to maintain shoot and root growth. A
negative correlation between these two hormones was observed only in the
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root system of salt stressed, hydroponically grown, plants (Picarella et al.
2007).
Scatolone di Bolsena
see Landrace, tomato
Senescence
see Programmed cell death
Spodoptera littoralis
Stefano Speranza, Gianluca Petti, Claudio Pucci (Tuscia University, Viterbo Italy)
Considering the biological relationship between anthocyanin (see
Anthocyanins) induction and insect attack and the availability of tomato
lines differing for anthocyanin content in the vegetative tissues of the plant,
experiments have been carried out in order to verify the effect of tomato
leaves with different anthocyanin content in the diet of Spodoptera littoralis
larvae, from the 4th living day to their pupating. To this aim, it has been
used a tomato line with high anthocyanin content in leaves and stems (line
V711088, somaclonal variant atvsc), a near isogenic line without
anthocyanins (line V711232, somaclonal anthocyaninless variant) and a
normal control line (V710040,
cv UC-82).
The data obtained during the
trial in temperature-controlled
environment
(see
Figure)
showed no differences during
the exponential growth of the
three cohorts. A first difference
is seen when the weight peak of
the larvae fed with high and zero
Weels with tomato leaves feeding
anthocyanin diet is reached two
Spodoptera larvae.
days in advance than the control.
Differences detected among the three cohorts for larvae survival were not
significant. These results suggest that the insect detects, at midgut level,
changes in the chemical composition of the leaves, both as an increase of
anthocyanins or as an absence, but that such condition is not sufficient to
induce death. However, the high anthocyanin content, or anthocyanin
absence, although not letal for Spodoptera larvae, could give sub-letal
effects to their physiology.
spongy seed (ss)
see Tomato, genetic markers
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stamenless-2 (sl-2)
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
We addressed the study of the stamenless-2 mutant both as a source of male
sterility for hybrid seed production (Badino 1976; Badino and Soressi
1976) and to test the relationship with the pat mutation (see parthenocarpic
fruit) because aberration of the anther cone
has been associated with parthenocarpy in
tomato and in other species. The two loci
were not allelic and the pat sl-2 double
mutant showed mainly epistatic effects; sl2 was epistatic to pat for phenotypes
related to male reproductive development
(anther phenotype and seed production
under open pollination), whereas the pat
mutation was the limiting factor for aspects
related to the female side (ovule phenotype,
An sl-2 mutant anther with
fruit size and seed production under hand
ectopic
ovules
and
pollination). Thus, the genetic basis for the
carpelloid structures.
floral defects in the two inner floral whorls
is different in the two mutants (Mazzucato et al. 2008b).
Genetic analysis indicated that the primary aberration of the sl-2 genotype
is the specification of stamen identity, supporting the hypothesis that the Sl
locus may correspond to the tomato ortholog of DEF. Although our work
supported this hypotesis with mapping information (both Sl and SlDEF
mapped on the long arm of chromosome 4, Mazzucato et al. 2008b), very
recent analysis of segregating progenies would indicate that the two loci are
not actually allelic.
sticky peel (pe)
see “Fruit skin” mutants
Sundwarf mutants
Agostino Falavigna (CRA – Research Unit for Vegetable Crops, Montanaso
Lombardo, Lodi Italy)
Franco Todisco (Università Cattolica del Sacro Cuore di Piacenza, Italy)
Searching for a tomato plant with reduced, compact habit of possible
advantage for open field cultivation and mechanical harvesting, Soressi’s
attention was drawn by the so called “bird nest” phenotype. The name
springs from the plant brachitism with curved leaves encircling the fruits
like in a nest. The first “bird nest” variety, with hp, sp, u genes, is “Ottawa
60” released in Canada in 1964 by Loyd Lyall.
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By considering the existence of the “sundwarf” (sd) mutant (described by
Rick in 1956) reduced in plant size, with internode progressively reduced in
relation to light intensity and soil conditions, and its phenocopies, induced
by
EMS
and
γ-rays
respectively in S. Marzano
and MoneyMaker, a series of
crosses involving sundwarf
mutants, two birdnest types
and lines and determinate (sp)
and indeterminate (sp+)
cultivars was planned. The
Fig.1 Sundwarf tomato plants with
extremely reduced habit (rosette).
numerous F2 progenies were
grown in open field for a not
easy and time consuming classification into distinct phenotypic groups
(Todisco 1978).
Notwithstanding the wide variation in plant habits ranging from
indeterminate normal type to a rosette-like
one (Fig. 1), the gathered data revealed the
existence of two sundwarf genes (sd-2,
incompletely
recessive,
and
sd-3),
interacting each other and with the sd, sp
and br Mendelian factors. In addition, it was
possible to recognize the birdnest
phenotypes of cv. Ottawa 60 and H-105
from Canada as due to sd-2, sp and br
homozygous genes (Falavigna and Soressi
1975, XIX SIGA Annual Congress;
Falavigna and Soressi 1978). The field trial
Fig.2 Well performing
carried out with a sundwarf “cluster”
sundwarf “cluster” tomato
phenotype (Fig. 2) at a population density of
plant in open field.
130,000 plants/hectar evidenced a yield
potential not significantly different from the best processing cultivars but
earlier in fruit ripening (Soressi 1974; Falavigna and Soressi 1975; Soressi
1979). In later work at the Tuscia University in 2001, the pro5::uidA
chimaeric reporter construct (see Lateral suppressor) was introduced in a
sundwarf line for future studies on the IAA metabolism in relation to the
plant habit influenced by different light spectrum and intensity.
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Tetranychus urticae
Marisa Castagnoli CRA-ABP (ex Istituto Sperimentale per la Zoologia Agraria)
Riccardo Caccia (Università della Tuscia, Viterbo Italy – presently different
address)
A critical aspect dealing with the use of transgenic plants is the global
evaluation of their environmental impact. Tomato, because an economical
important crop, and Tetranychus urticae (Acari Tetranychidae), a very
injurious polyphagous mite, are the case study chosen to investigate
unpredictable and undesirable effects on phytophagous arthropods. Three
tomato near isogenic lines, i.e. the cv. Riogrande (RIG) and the transgenic
lines RC332 [containing the Gox gene (see Glucose oxidase) and showing
high glucose oxidase activity], and MS498 [containing the KTI3 gene (see
Helicoverpa armigera) and exhibiting a high trypsin inhibition] were
considered. Trichomes and contents of C and N of the leaves, differences in
development and oviposition of T. urticae and damage caused were
evaluated for each line. Laboratory trials evidenced that 1) the intrinsic rate
of increase of two strains of T. urticae (T tomato-adapted strain, B nontomato adapted strain), was significantly lower in RIG than in transgenic
lines and doubling time ranged between 6.9 and 11.6 days in the first and
between 3.9 and 5.3 days in the latter; 2) the glandular four-lobed trichomes
were always higher in RIG than in other genotypes, 3) the N leaf content
was from 1.3 to 1.9 fold lower and the C/N ratio from 1.3 to 1.9 fold higher
in RIG than in other lines. Greenhouse experiments performed with an
initially equal infestation of strain T evidenced: 1) no significant difference
between plant lines in the final mite infestation (motile stages per plant),
nevertheless an almost double number of spider mites was counted in
RC332; 2) a significantly higher percentage of damaged leaves and a
significantly higher average damage index on RC332 than on RIG (79%
and 2.3 in the former, and 62% and 2.1 in the latter, respectively), even if in
both transgenics a higher level of the most severe damages and a shorter
time to approach them were observed; 4) a comparable number of mites
causing the same damage level in all genotypes and a strong linear
relationship between the first four levels of damage and mite infestation.
Although in the laboratory studies both transgenic lines enhanced the T.
urticae population increase, the glasshouse studies were not as conclusive
and they only suggested the possibility of real differences between the
transgenic and non-transgenic genotypes (Castagnoli et al. 2003).
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Tomato, “fruit colour genes”
Andrea Mazzucato, Maurizio Enea Picarella, Pietro Mosconi (Università della
Tuscia, Viterbo Italy)
The collection of a wide variety of germplasm entries and the selection of
new mutant types and alleles allowed to work at the breeding of materials
characterized by different colours of the fruit epidermis and flesh and, as
such, by different contents of nutritional compounds. This included mutants
and genes with altered composition of carotenoids (red, Beta, Delta),
anthocyanins (Aubergine, Anthocyanin fruit, atroviolaceum) and
chlorophyll (uniform, green flesh), with increased pigment composition
(High pigment-1 and –2, Intense pigmentation) or lacking a fraction of
pigments (colorless epidermis). A very first mention about the potential of
this mutations for diversifying the appearance and the composition of the
tomato products was given in early contributions (Soressi 1969b; 1969c).
Later, using the germplasm retrieved in foreign collections from Butler and
Darby, Soressi started to introgress genes and mutations for “colour genes”
in the most important genetic backgrounds at that time (Marmande, San
Marzano, Roma, New Yorker and Gimar). Such set of near isogenic lines
has been partially lost; only the Gimar and San Marzano series remain
active in Soressi’s collection. This breeding lines served as the basis for
research aimed at establishing the pattern of pigment composition and
red, high-pigment-2 and sun black (sb); the last is a recent result of Soressi’s
breeding with “color genes”.
antioxidant activity in different backgrounds (Minoggio et al. 2003)
Later, the study of tomato nutritional compounds has been undertaken using
a comparison of the spectral characterisation of the tomato fruit surface
pigments from the immature to over-ripe stage, using spectroscopy
techniques based on visible fluorescence emission upon excitation in the
same or ultraviolet spectral regions with the aim of verifying the spectral
band for optimal conditions for fruit harvesting using non-destructive
techniques. The main fluorescence spectral features belonging to
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anthocyanins, flavonoids, carotenoids and chlorophyll a after excitation of
skin tomato pigments at different laser wavelengths was identified (Lai et
al. 2007).
Another study investigated the combined effect of diluted seawater and
ripening on the beneficial nutritional properties of tomato fruits from an
antioxidant point of view. The research was conducted on tomato cv. Jama,
Gimar wild type, Gimar gf, and Gimar nor. The genotypes Jama and Gimar
gf showed increases in ascorbic acid, lipoic acid, and R-tocopherol during
both ripening and salt treatment, whereas total ascorbate and tocopherols
decreased in the berries from salt-treated plants of Gimar wild type.
Ripening also determined decreases in ascorbate and tocopherol amounts in
the Gimar nor line where a positive effect of ripening and salinity was
observed (Sgherri et al. 2007).
The research activity on “colour genes” has yielded collaborations with
private companies (Barilla SpA, Tomato Colors) and initiatives for
divulgation as a “bruschetta party” made with tomatoes of different colours
(Mazzucato et al. 2007; Mazzucato and Picarella 2007).
Tomato, general
Andrea Mazzucato, Maurizio Enea Picarella, Pietro Mosconi (Università della
Tuscia, Viterbo Italy)
The long and wide research activity resulted in a deal of review articles and
book chapters or books focussed on the tomato crop (Soressi 1969a; 1969b;
1974; 1976; 1979; 1992; Porcelli et al. 1979; Amadei et al. 1990; Acciarri
et al. 1998; Monti et al. 2004).
Such research activity has allowed to put together a collection of tomato
germplasm listing more than 650 accessions, that includes landraces,
vintage and modern cultivars, genetic stocks and several other materials
that, during the years, have undergone considerable characterisation.
Compatibly with the absence of dedicated resouces, the collection has been
maintained active, used for the internal research activity and distributed
worldwide. Recently the collection has been transferred to a database in
Microsoft Access format and more than 500 accessions have recently been
included in the ECPGR Tomato Database
(http://documents.plant.wur.nl/cgn/pgr/tomato/).
The more recent activity of the Soressi and Mazzucato group at the Tuscia
University has been described in a dedicated web site
(http://www.unitus.it/tomato/).
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Tomato, genetic markers
Andrea Mazzucato (Università della Tuscia, Viterbo Italy)
The use of natural or induced morphological variants, recognizable at an
early stage of plant development, has been proposed as an aid to the
production of hybrid tomato seeds under three different strategies (Soressi
1966; 1977; Sozzi 1969).
Visible markers linked to male sterility genes were proposed in the
perspective of maintaining genic male sterile parental lines, through early
recognition of male sterile segregant; this is the case of anthocyanin absent
(aa) and white virescent (wv) linked to male sterile-10 (ms-10, alias ms-35)
and of anthocyianin without (aw) liked to ms-15 (Soressi 1974).
Genetic markers expressed in the seed have been proposed to distinguish
and/or select seeds from selfing or natural (NCP) or hand-outcrossing. Such
markers may involve the seed
colour, such as brown seed
inherited as an endosperm trait
(bs, Soressi 1967a; Bin and
Soressi, 1973; Soressi et al. 1975)
or the seed coat structure, such as
spongy seed inherited as a
maternal trait (ss, Soressi 1967b)
or both (Soressi 1968). Further,
schemes using two, non allelic
mutations for the bs character
A Soressi’s scheme for hybrid seed
have been described (Soressi
production aided by two, non allelic
1970b).
mutations for the bs character
The bs marker, together with those
(Soressi 1970b).
expressed in the plantlet, such as
potato leaf (c), may also serve in assessing unwanted crosses in the
multiplication of seed parent lines with incomplete or conditional male
sterility.
Tomato, high temperature tolerance
Venkata Rami Reddy Sanampudi (Università della Tuscia, Viterbo Italy)
The Intergovernmental panel on climate change (IPCC, 2001) announced
that the global atmospheric temperature may raise 2.6°C by 2050 relative to
1990 and 5.8°C by 2010.Atmospheric temperature is the main important
factor determining plant growth and productivity, hence many researchers
are investigating high temperature effects on plant productivity. Alteration
of the unsaturation levels of chloroplast membrane fatty acids could be a
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useful method for enhancing the tolerance of plants to environmental
stresses especially temperatures (Somerville and Browse 1991). Transgenic
tobacco plants low in trienoic fatty acids exhibited high temperature
tolerance (Murakami et al. 2000).
Plastid omega-3 fatty acid desaturase
catalyzes the conversion of the dienoic fatty
acids to trienoic fatty acids in glycerolipids
which are the main components of
chloroplast membranes of plants. At the
University of Tuscia, we produced
transgenic tomato plants that express
double-stranded RNA (dsRNA) of tomato
plastid omega-3 fatty acid desaturase gene
Transformed tomato plant
which results in silencing of plastid omegaat flowering stage.
3
fatty
acid
desaturase
gene.
Transformation was confirmed by PCR. Now we got first generation of
transformed plants and we are going to test against high temperature and
membrane fatty acids analysis.
Tomato hybrids, bs-marker based
Giò Batta Bruzzone, Pasquale Martiniello, Agostino Falavigna (CRA – Research
Unit for Vegetable Crops, Montanaso Lombardo, Lodi Italy)
By omitting the use of the male sterile genes (see stamenless-2; positional
Fig.1 A bombus,
one of the most
NCP
responsible
insect in tomato.
Fig.2 Simple vibrating
device for improving
pollen recovery from
male parent plants.
sterile-2) we report briefly,
the research activity, carried out under Soressi’s
guide, focused on the practical utilization of the bs
seed marker, in male fertile seed parent lines, which
allows to directly recognize the F1 straw coloured
seeds (+/bs) among the brown (bs/bs) ones (see
Genetic markers).
etic
Fig.3 Plant of
the “Brown-1”
tomato hybrid
obtained with
bs/bs
seed
parent.
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This knowledge has led to the development of theoretical schemes aimed at
exploiting the Natural Cross Pollination (NCP) by pronubous insects (Fig.
1) and an optico-mechanical separation of the differently coloured seeds.
Due to the lack of an efficient and economically convenient electronic
equipment for F1 seed separation, Soressi’s attention was drawn on the
technique based on hand pollination of bs/bs autofertile seed parent lines
without flower emasculation (Bruzzone 1980; Falavigna et al. 1981). Such
a procedure allows to obtain 99-100% F1 seeds in relation with stigma
receptivity and pollen amount (Fig. 2). The synthesis of tomato hybrids
(+/bs) of agronomic and commercial value requires a skilfull work for
incorporating, through cross and backcross, the bs trait into the seed parent
line that shall be combined with a pollen donor with normal seed colour. To
choose the two parent lines, a previous work for evaluating the General and
Specific Combining Ability, in relation with the final product (table or
processing tomato), needs to be carried out (Maggiore et al. 1976;
Martiniello and Soressi 1977). As the bs is present in homozygous situation
in the seed parent lines and in heterozygous one in the F1 hybrids, the
possible marker’s effect on plant and fruit characteristics was ascertained.
Notwithstanding the delay in germination, initial seedling growth,
flowering and fruit ripening of the bs/bs lines, no drawback is expected
from using the bs gene, as marker, in the seed parent . In the 1977-79
triennium, a high number of F1 hybrids obtained by hand pollinating bs and
bs-2 seed parent lines were included and evaluated in an outdoor
comparison trials located in Northern, Central and Southern Italy. One of
the best of these hybrids was inscribed, in 1982, in the National Register of
Varieties with the name “Brown-1” (Fig. 3).
Tomato, hybrids for mechanical harvesting
Carlo Lorenzoni (Università Cattolica del Sacro Cuore, Piacenza Italy)
Tommaso Maggiore (Università di Milano, Italy)
The possibility of the practical exploitation of hybrid vigour in processing
tomato was studied in a period (early ‘70s) when attention at this regard
was given mostly to determinate growth types. Two series of hybrids were
produced, following a NC2 mating scheme (common mothers-common
fathers) with 106 x 4 and 86 x 3 entries, respectively, where nearly all the
parents were short vined (homozygous for the self pruning, sp, allele)
varieties. The data, collected in two locations (Northern and Southern
Italy), allowed an appreciation of the general combining ability of the single
varieties for the characters under observation, i.e. flowering and first
ripening time, time of picking, total and marketable yield, number of
marketable fruits, percent of scrapped fruits, average fruit weight, ratio
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between fruit diameters, degree of sun scald, degree of cracking, fruit
firmness, soluble solid content (Maggiore et al. 1973; Soressi et al. 1973;
Salamini et al. 1975; Maggiore et al. 1976; Conti et al. 1991).
A comprehensive evaluation of the results presented and discussed let also
draw some general conclusions:
- the hybrids are usually earlier than the varieties;
- as for the mean yield, the hybrids exceed the varieties by 20%, even
though in the best environment some varieties can reach the highest level of
production; when cultivated in suboptimal conditions, the F1s always
overcome the varieties;
- the hybrids seem to possess a higher phenotypic stability than the
varieties;
- the hybrids are clearly superior with regard to the fruit number per plant,
but their fruit size is lower; it must be emphasized that high yielding plants
are characterized by small or intermediate fruit size;
-the fruit quality is not influenced by the hybrid vigour.
High yielding F1s can be developed, by exploiting the genotypes explored:
these hybrids should bear 40-60 fruits possessing a length/width ratio of
1.2, a weight of 50-60 g, and good fruit firmness.
In a preliminary phase of the research, the efficiency in hybrid seed
production was investigated in relation to variables such as plant genotype,
location, frequency of pollinating insects, planting design, natural and hand
pollination with and without emasculation. The results suggested
procedures, combining genetic agronomic and entomological factors able to
increase natural cross pollination rates avoiding hand work (Soressi and
Lorenzoni 1970; Bin and Soressi 1973). Also suggestions for the use of
male sterilty were thoroughly considered (Badino and Soressi 1976, see
Male sterility; see stamenless-2).
Tomato, “high-pigment genes”
Maurizio Enea Picarella, Enrico Santangelo (Università della Tuscia, Viterbo Italy)
Antonella Lai (ENEA- Frascati, Roma Italy)
Alberto Pardossi (Università di Pisa, Italy)
In the middle of the 70’s, within Soressi’s breeding activity on fruit colour
genes to be introduced in different backgrounds, special attention was
driven to some mutants collectively referred to as “high-pigment group”. In
particular, high pigment (hp), high pigment-2 (hp-2) and green flesh (gf)
BC1S4 San Marzano breeding lines were studied. Whilst hp and gf had been
provided by prof. Leonard Butler (see Tomato, “fruit colour genes”), hp-2,
more recently isolated by C. Bowler (Mustilli et al. 1999), had been
induced by EMS seed treatment in cv. Garim, a S. Marzano type (Soressi
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78
1975). These monomendelian, recessive genes were attractive for
enhancing leaf and fruit pigments as proved by spectrophotometric and
HPLC measurements of chlorophyll a and b and carotenoids (Palmieri et al.
1975; 1978). To better clarify the effects of the “hp genes” on the
mechanism of pigment production in leaf and in fruit ripening, their
antioxidant value, chlorophyll a and b, total carotenoids and anthocyanins
content were determined by spectrophotometric analyses. The assessed
relationship between the
pigment content in the leaf
and the fruit at different
ripening stages led to the
opportunity of adopting a
more reliable method of
a
b
c
analysis (Santangelo et al.
Fig.1. Different fruit pigmentation at the
2002). For this reason total
breaking stage of S. Marzano NILs
polyphenols,
flavonols,
homozygous respectively for gf (a), gf, l-2 (b)
flavonones, phenolic acid,
and l-2 (c).
lycopene and β-carotene
were determined in different tomato mutant lines and cultivars through
mass-spectrometry and HPLC analyses; total antioxidant activity was also
measured. Opposite to a reasonable
increase of carotenoids in the fruit
of hp and hp-2, the level of
flavonoids and other phenolics was
lower than in the WT; reversely, it
increased in mutants [lutescent-2 (l2), rin/+, u] defective in
carotenoids accumulation in the
1
2
fruit (Minoggio et al. 2003). The
biological activities of polyphenols
are well known and today they are
more
appreciated
for
their
beneficial effects on human health.
Thus, when defining the total
antioxidant activity of fruit and
4
3
vegetables, polyphenols, besides
Fig.2. Macroscopic effect of
carotenoid concentrations, should
continuous light stress on tomato
be considered as well.
leaves of green flesh (1), lutescentBecause in the gf mature fruit some
2 (2), double recessive (gf, l-2) (3)
of
the chloroplasts are not
and WT (4) NILs.
transformed into chromoplasts, this
mutant was judged particularly suitable in revealing possible levels of lipoic
acid (LA), one of the most efficient biological antioxidants that mostly
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occurrs in green plant organs. In relation also to this issue, the capacity of
salinity stress to increase the nutritional value of tomato (see Salt
tolerance), was investigated in the commercial table tomato F1 hybrid Jama
and in Gimar NILs (Gimar-WT, Gimar-nor, and Gimar-gf) grown in
closed-loop rockwool culture. A striking result was the increase of the
antioxidants ascorbic acid, tocopherols (α and γ), and to a minor extent of
LA, in Gimar-gf fruits in comparison to Gimar-WT and Jama when plants
were exposed to moderate salinity conditions (Sgherri et al. 2007).
Now, it is known that hp-1 and hp-2 are isophenotypic mutations with
exaggerated light responsiveness. The Hp-1 locus is the tomato ortholog of
the gene encoding UV-damaged DNA Binding Protein 1 (DB1) and the
phenotype of hp-2 is caused by alteration of DEETIOLATED-1 tomato
ortholog. Both are responsible for enhancing the expression of genes
involved in the production of fruit pigments via an interaction with
chromatin. The gf mutation displays a non-functional stay-green phenotype
with delayed chlorophyll degradation in the leaf during senescence and in
the fruit during ripening. Efforts are being put forward in the Laboratory of
vegetable biotechnology at the Tuscia University aiming to identify the
lesion responsible for the gf phenotype. These are carried on by tools such
as genetic analysis, microsynteny, physical mapping, functional analysis of
candidate genes. Of particular interest has appeared the combination of gf
with l-2, a mutant accelerating chlorophyll degradation. In double recessive
plants, the presence of gf reduces the progressive de-pigmentation
(yellowing) of the green parts, well noticeable in both l-2 leaf and fruit (Fig.
1).
Such kind of interaction works also at the physiological level in a beneficial
way for plant stress response such as continuous light exposure (Fig. 2).
Physiological analyses on mutant lines with a role in plant senescence, like
lesion mimic, and in fruit pigmentation enhancing (hp-1, hp-2), combined
with gf and l-2, are also in the pipeline.
Tomato, leaf peroxidases
Gian Piero Soressi (Università della Tuscia, Viterbo Italy)
The leaf peroxidase activity and the electrophoretic banding pattern of 69
tomato mutants affecting plant morphology have been studied (Soressi et al.
1974). The zymograms of 63 mutants were normal, whereas six mutants
(lyrate, mottled, olivacea, monstruosa, extreme dwarf and Curl) were
characterized by one or two more intensely stained bands. Their total
peroxidase activity was significantly increased reaching the maximum in
olivacea (40-folds higher than the WT). It could be inferred that the six
abovementioned mutations influence directly or indirectly the peroxidase
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activity level without affecting the genes which code for the peroxidase
isozymes themselves. Parallels between the mutant effects on specific
peroxidase bands and hormonally-mediated control of the peroxidase
isozymes are discussed. Later, the activity of IAA oxidase in the mutants
monstruosa and olivacea were studied (Palmieri et al. 1978). Analyses of
the inheritance of the peroxidase activity and relative electrophoretic
isoenzyme bands were olds done in crosses and backcrosses involving L.
esculentum mutants and cvs with L. hirsutum accessions (Giovinazzi 1978).
More recently, peroxidase activity was studied as a player in cell death
responses controlled by the Pto-Fen complex following fenthion treatment
(De Biasi et al. 2003, see Fen gene).
Triploidy
see Polyploidy
uniform (u)
see Tomato, “high-pigment genes”
Waste heat utilization in agriculture
Massimo Schiavi (CRA- Research Unit for Vegetable Crops, Montanaso Lombardo,
Lodi Italy)
Tommaso Maggiore (Università di Milano, Italy)
When Soressi was head of the ISPORT Montanaso Lombardo (Lodi) out
Section, CTN-ENEL, Milan planned (1977) and built (1978-1980) an
experimental agro-thermic
pilot plant at TavazzanoMontanaso power station
(Fig. 1) availing itself of the
scientific and technical
collaboration of the ISPORT
Section Staff, in this case
under M. Schiavi main
responsibility and G.P.
Soressi and T. Maggiore
coordination.
The
experimental activity, aimed
Fig.1 Overview of the experimental agrothermic pilot plant at Tavazzano-Montanaso
to prove technical and
power station.
economic feasibility of
using waste water heat
energy from the abovementioned station for crop production, was
developed in two phases: an optimization term for identifying technical
parameters to exploit at the highest level the relatively low (∆t=8°-9°C)
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thermic gradient available; a second term for managing the crop cultivation
trials for achieving the yield and related economic parameters.
To obtain earlier crop production without negatively affecting marketable
yield quantity and quality, the following experiments in plastic tunnels or
open air were scheduled in relation to the different tested crops (Soressi et
al. 1981):
• Asparagus (green and white spears): in 3 tunnels with “EVA” plastic
cover; 1 unheated and 2 heated with waste water in 4 polyethylene
(PE) pipes (2 buried in the soil under the crowns and 2 over soil (8-10
cm) in open air plot, unheated or heated in the same way.
• Strawberry: “EVA tunnels” (1 unheated, 1 heated in both soil and air).
• Melon: “EVA tunnels” (1 unheated, 1 heated soil and air).
• Lettuce: for 3 culture cycles (September-April) in 3 glasshouses, 1
unheated, 1 heated in both soil and air and 1 heated in soil, air and
with heat water running on the roofs.
• Maize: late-ripening pure lines to be used as parents in F1 hybrid seed
production, to get earlier yield seeds (heated and unheated big-plot in
open air).
• Germinating cereal seedlings: barley, maize and wheat for integrating
diary animal forage.
The best results putting in value the energy waste were obtained with maize
pure lines, asparagus and strawberry. Regarding maize, the flowering was
5-8 days anticipated with a yield increment (up to 80%); in asparagus a
significant earliness in spear emission and yield in both tunnels (26 days)
and open air (18 days) was reached; as to the strawberry, the latest varieties
gave an earlier (20 days) yield
increase (11%) (Schiavi et al.
1983; Schiavi et al. 1984b;
1984c; Rotino et al. 1985).
By adopting a heat pump in
1983/84, a new trial for
evaluating yield of Gerbera cutflowers was set up in two glasshouses where the cv. Boston
performed as the best in the
autumn-winter period (Fig. 2)
Fig.2 Gerbera cultivation in glasshouse,
(Schiavi et al. 1984a). The
aerothermically heated through 20°C
possible advantages connected
water circulation.
with the utilization of the water
low thermal level from electric power-plant are strictly dependent on the
economic convenience of some technical actions aiming to avoid heat loss,
rationalize the heat flux in relation to luminosity variations and to the
cultivation techniques. Parallely, an aid system is recommended because
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the needed continuous availability of hot water is not guaranteed being
dependent on the working cycles of the electric power-plant (Soressi et al.
1985).
In the subsequent decennium (1986-96) such a synergistic cooperation,
between ISPORT section and CTN-ENEL at Tavazzano-Montanaso electric
power plant, prosecuted by carrying out trials with Gerbera, leafy
ornamentals, pepper, eggplant and tomato by adopting different soilless
culture techniques.
Watermelon, germplasm evaluation
Carlo Lorenzoni (Università Cattolica del Sacro Cuore, Piacenza Italy)
Basilio Borghi (Parco Tecnologico Padano, Lodi Italy)
At the beginning of the years ‘70s the observation that new cultivars of
watermelon (Citrullus vulgaris L. Schrad) of foreign origin were
substituting the local varieties on the Italian market, evidenced the need of
an evaluation of the traditional germplasm as a source for breeding
programmes. A collection of 43 cvs coming from Italy, Japan, Russia and
USA was explored in order to: 1) obtain information about the inheritance
of a series of characters (earliness of flowering and ripening; fruits per
plant; firmness of rind; rind thickness / transversal diameter of the fruit
ratio; sugar content of the ripe fruit juice; number and weight of the seeds
per fruit; 1000 seeds weight); 2) evaluate the single cultivars as source of
useful traits for the breeding; 3) estimate the possible advantages of the
hybrids in comparison with the varieties; 4) put the basis for future breeding
programmes (Bergamaschi 1971).
The varieties and 22 hybrids, having at least one Italian variety as a parent,
were compared in the field. In the trials, Italian varieties appeared
outstanding as donors of useful characters for fruit setting and total yield,
likely connected with a good environmental adaptation; USA entries were
notable for yield and sugar content; the Japanese ones emerged for
earliness, sugar content and low seed number; both Italian and Russian cvs
resulted good sources for rind firmness.
Crosses between Italian and imported genotypes seemed highly promising
for selection in the segregating generation. On the contrary, the hybrids per
se rarely demonstrated of value for a direct exploitation in cultivation. That
does not exclude the possibility of individuating cross combinations
convenient for the crop, but likely specific and wider programmes are
required in order to find them (Soressi et al. 1974).
For hybrid seed production technique in the cucurbitaceae, also the use of
Ethrel as a phytoregulator able to suppress male flowers was evaluated
(Borghi et al. 1972).
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waxy fruit (pe-2)
see “Fruit skin” mutants
Xanthomonas (resistance to)
see “Lesion mimic” mutants
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BIBLIOGRAPHY
Acciarri N, Di Candilo M, Sanguineti MC, Soressi GP, (1998) Tomato. In: Italian
Contribution to Plant Genetics and Breeding, G.T. Scarascia Mugnozza & M.A.
Pagnotta (eds.), pp. 463-465. University of Tuscia, Viterbo
Allavena A (1975) Induzione di forme polipoidi in L. esculentum Mill. Omozigote per il
fattore pat di partenocarpia. Graduation thesis, Università Cattolica Sacro Cuore,
Piacenza, Italy. Tutor Prof. GP Soressi
Allavena A, Falavigna A, Soressi GP (1976) Forme polipoidi partenocarpiche
artificialmente indotte in Lycopersicon esculentum Mill. Colture Protette V(8-9):
45-46
Allavena A, Soressi GP (1978) Tetraploid parthenocarpic fruitful tomatoes. Tomato Gen
Coop Rep 28: 2
Allavena A, Lancelotti M, Palmieri S, Soressi GP (1978) Influenza del gene nor2 su
alcune caratteristiche qualitative della bacca di pomodoro durante la maturazione e
la post-maturazione della pianta. I Conv. Naz. “Problemi e prospettive del
pomodoro da mensa”, Monsampolo del Tronto, 7-8/11/1978 I.SP.ORT., Salerno,
145-164
Allavena A, Soressi GP (1982) Inclusione di tolleranza e/o resistenza alla batteriosi ad
alone ed al virus del mosaico comune in fagiolo da granella (Phaseolus vulgaris
L.). La Difesa delle Piante 5-6: 405-41
Allavena A, Fadda A, Soressi GP (1982a) New dry bean (Phaseolus vulgaris L.)
varieties resistano to BCMV. Genet. Agr. 37: 138-139
Allavena A, Del Zan F, Fadda A, Murgut G, Siviero P, Tonetti T, Soressi GP (1982b)
Attività di costituzione di varietà di fagiolo nano resistenti a BCMV e a
Pseudomonas phaseolicola. Informatore Agrario 38 : 20625-20639
Allavena A, Fadda A, Soressi GP (1985a) "Niveo” e “Montebianco”: varietà di fagiolo
(Phaseolus vulgaris L.) a seme bianco resistenti al virus del mosaico comune
(BCMV). Sementi Elette, 31: 39-40
Allavena A, Fadda A, Soressi GP (1985b) “Montalbano” e “Mogano” varietà di fagiolo
da granella (Phaseolus vulgaris L.) resistenti al virus del mosaico comune ottenute
a seguito di trattamenti con MSE. Sementi Elette, 31: 37-38
Allavena A, Fadda A, Soressi GP, Salamini F (1986) Grazia and Patrizia bush bean.
Hortscience 2: 1081-1082
Allavena A, Fadda A, Soressi GP (1989) “Montecarlo” and “Monterosa”, “Borlotto”type dry beans. Hortscience 24: 1047-48
Allavena A, Angelini R, Genga AM, Pernacchia G and Soressi GP (1994) Advanced
bean tissue culture and transformation research in Italy. Proceedings BARN
NETWORK- CIAT, Cali, Colombia, 7-10 Settembre, 1993: pp 190-198
Amadei G, Trentini L, Soressi GP (1990) Il pomodoro – The tomato. Enichem
agricoltura e Agrimont, Ed. unica Milano pp X + 91
Angelini Rota R (1987) Morfogenesi in vitro da tessuti somatici di species del genere
Phaseolus. Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor Prof. GP Soressi
Antognoni F, Ghetti F, Mazzucato A, Franceschetti M, Bagni N (2002) Polyamine
pattern during flower development in the parthenocarpic fruit (pat) mutant of
tomato. Physiol Plant 116: 539-547
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
85
Antonelli MM, Ferrari G, Portesi A, Fogher C, Soressi GP (1987) Plant regeneration in
L. esculentum UC-82 and torosa-2 genotypes. Eucarpia Tomato Working Group, 26 September 1987, Salerno-ISPORT.
Antonelli M, Di Baccio D, Ederli L, Francini A, Marabottini R, Pellegrini E, Ciaffi M,
Lorenzini G, Nali C, Pasqualini S, Santangelo E, Sebastiani L, Soressi GP, Badiani
M (2005) Ozone as a tool for studying stress responses in plants: signalling and
defence in normal, mutant and transgenic tomato lines. Proceedings XV Meeting of
the Eucarpia Tomato Working Group, Bari, 20-23 September 2005. In press in
Acta Horticulturae
Antonelli M, Marabottini R, Santangelo E, Francini A, Pellegrini E, Di Baccio D, Ederli
L, Nali C, Pasqualini S, Soressi GP, Ciuffi M, Lorenzini G, Sebastiani L, Badiani
M (2007) Ozone as a tool for studying stress responses in tomato (Solanum
lycopersicum L.). III. Ethylene, cyanide and the development of foliar symptoms in
the autonecrotic mutant V20368. In: Ramina A et al. (eds.) Advances in Plant
Ethylene Research: Proceedings of 7th International Symposium on the Plant
Hormone Ethylene (Pisa, 18-22 June 2006): 389-390
Astolfi S, Zuchi S, Picarella ME, Passera C, Soressi GP (2005) Salinity induces changes
in nitrogen metabolism in tomato plants differing in salt tolerance. In: Plant
nutrition for food security, human healt and environmental protection. C.J. LLi et
al., (Eds.), 2005. Tsinghua University Press. Beijing, China; pp. 576-577
Badino M (1976) Il mutante di L. esculentum Mill. "short anthers": Eredità e sua
utilizzazione del breeding. Graduation thesis, Università Cattolica Sacro Cuore,
Piacenza, Italy. Tutor Prof. GP Soressi
Badino M, Soressi GP (1976) La maschiosterilità nella produzione di ibridi di pomodoro
per la serra. Colture Protette 5: 41-45
Balestra GM, Caccia R, Antonelli MG, Soressi GP, Varvaro L (2000) Population
dynamics of Xanthomonas visicatoria and Pseudomonas syringae pv. tomato on
tomato plants expressing specific (Pto) and aspecific (Gox) resistance genes. J
Plant Pathol 82: 77
Baroni D (1994) Confronto tra genotipi di pomodoro differenziati per portamento e
partenocarpia in coltura anticipata di piena campo. Graduation thesis, Università
Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Bedini C, Caccia R, Triggiani D, Mazzucato A, Soressi GP, Tiezzi A. Micropropagation
of Aloe arborescens Mill.; a step towards efficient production of its valuable leaf
extracts. Submitted to Plant Biosystems
Beraldi D (2001) Ricerca di marcatori molecolari associati al gene di partenocarpia
parthenocarpic fruit (pat) in pomodoro. Graduation thesis, Università della Tuscia,
Viterbo, Italy. Tutor Prof. A Mazzucato
Beraldi D, Picarella ME, Soressi GP, Mazzucato A (2004) Fine mapping of the
parthenocarpic fruit (pat) mutation in tomato. Theor Appl Genet 108: 209-216
Bergamaschi G (1971) Valutazione di cultivar di anguria e di loro incroci in funzione
del miglioramento genetico. Graduation thesis, Università Cattolica Sacro Cuore,
Piacenza, Italy. Tutor Prof. GP Soressi
Bianchi A, Marchesi G, Soressi GP (1963) Some results in radiogenetical experiments
with tomato varieties. Rad Bot 3: 333-343
Bianchi A, Marchesi G, Soressi GP (1964) Alcuni risultati in esperimenti di
radiogenetica con diverse cv. di pomodoro. Genetica Agraria 18: 317-318
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Bianchi A, Soressi GP (1969) Mutanti di pomodoro artificialmente indotti suscettibili di
utilizzazione nel miglioramento genetico. Sementi Elette XV (3), 2-6
Bianchi A, Borghi B, Lorenzoni C, Salamini F, Soressi GP (1973) Acquisizioni di
genetica vegetale e insegnamento. Atti Ass Genet Ital 18:79-90
Bima PJ, Mensurati F, Soressi GP (1993) Rigenerazione da ipocotile in pomodoro (L.
esculentum Mill.) su substrato privo di ormoni. Atti XXXVII Convegno Annuale
SIGA, Orvieto, 11-14 Ottobre 1993:184 (abstract)
Bima PJ (1997) Studio dei fattori che condizionano la micropropagazione in diverse
cultivar di aglio (Allium sativum L.). PhD Thesis, Università della Tuscia, Viterbo,
Italy. Tutor Prof. GP Soressi
Bima PJ, Mensurati F, Soressi GP (1995) Effects of gibberellic acid (GA3) and
micropropagation on axillary shoot induction in monostem genotype (to-2) of
tomato. Current Issues in Plant Molecular and Cellular Biology. M Terzi, R Cella,
A Falavigna (Eds). Kluwer Academic Publisher, London, pp 411-416
Bin F, Soressi GP (1973) Insetti pronubi e produzione di seme ibrido di pomodoro.
Genetica Agraria 27: 35-74
Blanco A, Soressi GP (1988) Ibridazione interspecifica ed intergenerica. In: GT
Scarascia Mugnozza (ed) Miglioramento genetico vegetale. Collana: Scienza e
tecniche delle produzioni vegetali - Patron, Bologna. pp. 298-308
Borghi B, Maggiore T, Soressi GP, Poma Treccani C (1972) Tecnica di produzione di
seme ibrido nelle cucurbitacee mediante l’impiego dell’Ethrel. Riv Orto Floro
Frutticoltura Italiana 56: 750-759
Bricchi D (1993) Espressione del gene pat-2 e variazione di alcuni regolatori endogeni
di crescita in due versioni quasi isogeniche – normale e partenocarpica – di un
ibrido di pomodoro allevato in tunnel freddo. Graduation thesis, Università
Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Broadway RM, Duffey SS (1986) Plant proteinase inhibitor: mechanism of action and
effect on the growth and digestive physiology of larval Heliothis zea and
Spodoptera exigua. J Insect Physiol 32: 827-833
Bruzzone GB (1980) Una nuova tecnica per la produzione su larga scala di seme ibrido
di pomodoro: impollinazione, senza emasculazione, di linee portaseme
maschiofertili geneticamente marcate. Graduation thesis, Università Cattolica Sacro
Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Bruzzone GB, Falavigna A, Soressi GP (1983) Metodologia per la sintesi di ibridi F1 di
pomodoro mediante impiego del marcatore bs del seme. Giornate GOF-SOI,
Ferrara, 9 December 1983, Informatore Agrario n.s.: 45-47.
Bruzzone G B, Carnevali G, Frangi P and Soressi GP (1984). Il pomodoro “monostelo”:
stato attuale della ricerca. Notiziario tecnico. ERSO, 17, 7-13.
Caccia R (1998) Impiego del transgene codificante l’enzima glucosio-ossidasi (GOD) in
pomodoro ai fini della comprensione e manipolazione della tolleranza a stress
biotici. PhD thesis. Università della Tuscia, Viterbo, Italy. Tutor Prof. GP Soressi.
Caccia R, Delledonne M, Balestra MG, Varvaro L, Soressi GP (1999) Plant-bacterial
pathogen interaction modified in transgenic tomato plants expressing the Gox gene
encoding glucose oxidase. In "Genetics and Breeding for Crop Quality and
Resistance”. Kluwer Academic Publisher, Netherlands, pp 119-125
Caccia R, Delledonne M, Levine A, De Pace C, Mazzucato A (2001) Apoptosis-like
DNA fragmentation in leaves and floral organs precedes their developmental
senescence. Plant Biosystems 135: 183-190
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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Cammareri G (2003). Rigenerazione e trasformazione del mutante lateral suppressor
(ls) con un gene artificiale di partenocarpia (DefH9-iaaM) ed incremento
dell’efficienza di micropropagazione in pomodoro (Lycopersicon esculentum
Mill.). Graduation thesis, Università della Tuscia, Viterbo, Italy. Tutors Proff. GP
Soressi and A Tiezzi
Campion B (1977). Potenzialità del mutante torosa-2 nel miglioramento genetico del
pomodoro per raccolta meccanica. Graduation thesis, Università Cattolica Sacro
Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Campion B, Fadda A, Soressi GP (1981) Conversione a taglia nana del fagiolo di
Spagna rampicante (P. coccineus L.). Proceedings of “Risultati di un quinquennio
di lavoro sul miglioramento genetico del fagiolo da granella” Meeting, University
of Turin, 3 July, Cuneo Italy
Campion B, Falavigna A, Soressi GP, Schiavi M (1985) Efforts for in vitro androgenesis
in onion (Allium cepa L.). Genet Agr 39: 305
Campion B (1995) ‘Venere’ and ‘Alarico’, New Scarlet Runner Bean (Phaseolus
coccineus L.) Cultivars with Determinate Growth Habit. HortScience 30: 14831484
Cantoni M (1991) Influenza del gene pat-2 su allegagione e caratteristiche produttive
delle versioni quasi isogeniche normale e partenocarpica di un ibrido F1 di
pomodoro allevato in tunnel freddo. Graduation thesis, Università Cattolica Sacro
Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Caporali F, Soressi GP (1993) Ruolo dell'orticoltura nella prospettiva di un sistema
agricolo integrato. Informatore Agrario 49 (7 suppl.): 86-88
Carrai C, Fioravanti L, Tomassini C, Serra G, Soressi GP (1990) Miglioramento
genetico della Dieffenbachia. I. Tecnica di incrocio. Colture Protette 11: 107-114
Carrai C, Fioravanti L, Cavallini A, Serra G, Soressi GP (1993) Caratterizzazione
citologica di cultivar di Dieffenbachia. Colture protette 2: 101- 104
Castagnoli M, Caccia R, Liguori M, Simoni S, Marinari S, Soressi GP (2003) Tomato
transgenic lines and Tetranychus urticae: changes in plant suitability and
susceptibility. Exp Appl Acarol 31:177-188
Castelnuovo M (1984) Studio di fattori che influenzano l’embriogenesi somatica e la
micropropagazione in vitro del fagiolo. Graduation thesis, Università Cattolica
Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Chiappari A (1993) Indagine sull’accumulo e la distribuzione della sostanza secca e
alcune sue componenti in genotipi monostelo e normale di pomodoro (L.
esculentum). Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor Prof. GP Soressi
Colla G, Rouphael Y, Cardarelli M, Mazzucato A, Olimpieri I (2007) Growth, yield and
reproduction of dwarf tomato grown under simulated microgravity conditions.
Plant Biosystems 141: 75-81
Conti S, Di Candilo M, Soressi GP, Sanguineti MC, Dadomo M, Del Re L, Pasotti PP
(1991) Indagini su ibridi di pomodoro da industria. Sementi Elette 37(1): 9-16
Cravedi P (1966) Mutanti artificiali indotti in cultivar di pomodoro. Graduation thesis,
Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Cravedi P, Soressi GP (1969) Mutagenic effects of EMS in tomato under different
treatment conditions. Genetica Agraria 23: 235
Crinò P, Cristinzio G, Gentile A, Haegi A, Rugini E, Soressi GP (1996) Filtrati colturali
e tossine nella selezione di piante resistenti a malattie. Petria 6 (suppl. 1):197-217
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TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
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Cunico C (1991) Caratterizzazione di varianti somaclonali indotti in pomodoro
(Lycopersicon esculentum) a seguito di rigenerazione in vitro. Graduation thesis,
Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
De Biasi MG, Astolfi S, Acampora A, Zuchi S, Fonzo V, Santangelo E, Caccia R,
Badiani M, Soressi GP (2003) A H2O2-forming peroxidase rather than a NAD(P)Hdependent O-2 synthase may be the major player in cell death responses controlled
by the Pto-Fen complex following Fenthion treatment. Funct Plant Biol 30: 409-17
De Logu G (1974) Partenocarpia in Lycopersicon esculentum Mill.: dati relativi a
progenie F3 omozigoti per un fattore monogenico inducente lo sviluppo di bacche
apirene. Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor
Prof. GP Soressi
Di Rollo B (1998) Trasformazione genetica del pomodoro mediante Agrobacterium
tumefaciens utilizzando il gene AtCyS di Arabidopsis codificante una cistatina per
indurre resistenza ad insetti dannosi. Graduation thesis, Università della Tuscia,
Viterbo, Italy. Tutor Prof. GP Soressi
Donati P (1990) Produzione di seme ibrido in cipolla mediante maschiosterilità
citoplasmatica. Uso di gametocidi e tecnica seed-to-seed con valutazione tecnicoeconomica. Graduation Thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor Prof. GP Soressi
Falavigna A and Soressi GP (1978) Birdnest phenotypes as related with sundwarf genes.
Tomato Genet Coop Rep 28: 6
Falavigna A, Badino M, Soressi GP (1978) Potential of the monomendelian factor pat in
the tomato breeding for industry. Genetica Agraria 32, 159-160 (Abstr.)
Falavigna A, Badino M, Bruzzone GB, Soressi GP (1981) A new technique for tomato
hybrid seed production: direct hand pollination of male firtile bs lines. Acta
Horticulturae 111: 47-54.
Falavigna A, Roversi N, Soressi GP (1982) Brown-1: ibrido F1 di pomodoro da
industria realizzato con linee portaseme bs. Informatore Agrario 20: 21017-18.
Falavigna A and Soressi GP (1980) Ibridi di pomodoro da industria: risultati conseguito
nel triennio 1977-79 e prospettive. Atti Convegno su Miglioramento Genetico del
pomodoro da industria, Sorrento Sant’Agnello, 8-9 November 1972. Arti Grafiche
Del
Falavigna A, Soressi GP (1981a) Miglioramento genetico dell’asparago di Altedo: stato
attuale delle ricerche. Seminario sulla coltura dell’asparago, Villa Salina (BO) 29
giugno 1979, pp. 9-15
Falavigna A, Soressi GP (1981b) La coltura in vitro di antere: una efficace tecnica per la
sintesi di linee pure in Asparagus officinalis L. Genet Agr 35: 62-64
Falavigna A, Tacconi MG, Soressi GP (1983) Recent progress in asparagus (Asparagus
officinalis L.) breeding by anther vitro culture. Acta Horticulturae 131: 215-222
Falavigna A, Casali PE, Mazzanti C, Soressi GP (1984a) Metodologia per la sintesi di
ibridi tra cloni eterozigoti di asparago. Atti giornate GOF-SOI, 9 December 1983,
Ferrara, Italy, pp. 20-21
Falavigna A, Tacconi MG, Casali PE, Soressi GP (1984b) Variability in asparagus
(Asparagus officinalis L.) following anther in vitro culture. Genet Agr 38: 330-331
Falavigna A, Soressi GP (1987) Influence of the pat-sha gene on plant and fruit traits in
tomato (L. esculentum Mill.). In: Modern trends in tomato genetics and breeding,
p. 128. Proceedings of the 10th Meeting of the EUCARPIA Tomato Working
Group, 2-6 Sept. 1987, Salerno, Italy
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
89
Falavigna A, Soressi GP (1993) Miglioramento genetico per la qualità di specie orticole.
Informatore Agrario 49 (6): 61-65
Falavigna A (2007) Asparago: resa e qualità col miglioramento genetico. Agricoltura
19: 85-87 (Regione Emilia Romagna)
Falavigna A, Huaisong W, Alberti P, Casali PE, Mennella G, Toppino L (2008)
Interspecific hybridization for asparagus breeding in Italy. Acta Horticulturae 776:
291-298
Fantino MG, Schiavi M (1987) Onion breeding for tolerance to Fusarium oxysporum f.
sp. cepae, in Italy. Phytopathol Medit 26: 108-112
Fioravanti L, Carrai C, Tomassini C, Serra G, Soressi GP (1990) Miglioramento
genetico della Dieffenbachia. II. Programmazione della fioritura mediante
trattamenti con GA3. Colture Protette 11: 103-107
Frangi P (1984) Confronto per parametri di sviluppo e di produzione tra genotipi di
pomodoro (L. esculentum Mill.) monostelo e normale. Graduation thesis, Università
Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Frangi P and Soressi GP (1986). Confronto per parametri di sviluppo tra genotipi
monostelo e normale di pomodoro (Lycopersicon esculentum Mill.). Annali Facoltà
di Agraria, Università Cattolica S. Cuore, Milano 26(1), 15-28.
Gandolfi G (1992) “Breeding lines” monostelo per la costituzione di varietà e ibridi di
pomodoro da industria. Graduation thesis, Università Cattolica Sacro Cuore,
Piacenza, Italy. Tutor Prof. GP Soressi
Gianfilippi F (2006) Studio della diversità molecolare in popolazioni locali italiane di
lenticchia (Lens culinaris Medik) tramite marcatori ISSR. Graduation thesis,
Università della Tuscia, Viterbo, Italy. Tutor Prof. A Mazzucato
Giovinazzi F (1978) Controllo dell’attività perossidasica e delle bande in incroci di
Lycopersicon esculentum Mill. e L. hirsutum E. Bompi. Graduation thesis,
Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Gorguet B, Schipper D, van Heusden AW, Lindhout P (2006) High-resolution Wne
mapping of ps-2, a mutated gene conferring functional male sterility in tomato due
to non-dehiscent anthers. Theor Appl Genet 113: 1437–1448
Gorguet B, Eggink PM, Ocaña J, Tiwari A, Schipper D, Finkers R, Visser RG, van
Heusden AW ( 2008) Mapping and characterization of novel parthenocarpy QTLs
in tomato. Theor Appl Genet 116:755-67
Habashy AA (1993) Miglioramento della tolleranza allo stress salino in pomodoro:
variabilità' somaclonale indotta a seguito di rigenerazione in presenza di NaCl ed
individuazione di parametri in vitro utili nella selezione. PhD Thesis, Università
della Tuscia, Viterbo, Italy. Tutor Prof. GP Soressi
Habashy AA, Testa G, Mosconi P, Caccia R, Mazzucato A, Santangelo E, Soressi GP
(2004) Parthenocarpy restores fruitfulness in sterile triploid (3x) tomatoes
artificially obtained by crossing 4x × 2x somaclones. J Hort Sci Biotechnol 79:
322-328
IPCC (2001) Climate change 2001:impacts,adaptation and vulnerability technical
summary (www.ipcc.ch)
Kagan-Zur V, Yaron-Miron D, Mizrahi Y (1991) A study of triploid tomato fruit
attributes. J Am Soc Hort Sci 116: 228-31
Kiferle C (1998) Induzione di poliploidi partenocarpici in pomodoro mediante
rigenerazione in vitro di espianti da ipocotile di genotipi pat e pat-2. Graduation
thesis, Università della Tuscia, Viterbo, Italy. Tutor Prof. GP Soressi
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
90
Lai A, Santangelo E, Soressi GP, Fantoni R (2007) Analysis of the main secondary
metabolites in tomato (Lycopersicon esculentum Mill.) epicarp tissue during fruit
ripening using fluorescence techniques. Postharvest Biol Technol 43: 335-342
Lancellotti M (1978) Effetti del gene “non ripening” (nor2) sulla qualità della bacca di
pomodoro durante la maturazione. Graduation thesis, Università Cattolica Sacro
Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Lioi L, Piergiovanni AR, Soressi GP, Nigro C, Tamietti G, Turina M, Campion B
(2007) Caratterizzazione, selezione, risanamento e valutazione di cultivar locali di
fagiolo comune (Phaseolus vulgaris L.). Italus Hortus 14: 31-40
Maggioni L, Soressi GP (1992) Risorse genetiche delle specie orticole e loro
salvaguardia. pp. 64-97. In "Ambiente Italia". Lega per l'Ambiente pp. 501.
Arancia blu. Vallecchi editore, Firenze
Maggiore T, Lorenzoni C, Soressi GP, Mariani BM, Salamini F (1972) Ibridi opaco-2 in
Italia settentrionale. Maydica 17: 67-94
Maggiore T, Borghi B, Lorenzoni C, Mariani BM, Salamini F, Soressi GP (1973)
Determinated hybrid tomatoes. I. Comparison between hybrids and varieties and
definition of a standard fruit type for mechanical harvesting. Genet Agr 27: 235280
Maggiore T, Motto M, Salamini F, Soressi GP (1976) Attitudine combinatoria per
l’adattabilità alla coltura protetta in pomodoro (L. esculentum Mill.). Sementi
Elette 22: 9-16
Mapelli S, Frova C, Torti G, Soressi GP (1978) Relationship between set, development
and activities of growth regulators in tomato fruits. Plant Cell Physiol. 19: 12811288
Mapelli S, Torti G, Badino M, Soressi GP (1979). Effects of GA3 on flowering and fruitset in a mutant of tomato. HortSci 14: 736-737
Mapelli S, Bricchi D, Cantoni M, Soressi GP (1995) Gene pat-2 e livelli di fitoregolatori
endogeni, allegagione e caratteristiche produttive in pomodoro. Italus Hortus 2:7479
Marchesi G, Soressi GP (1965) Risposta di due cultivar di pomodoro al trattamento con
metansulfonato di etile. Atti Ass Genet It Pavia 10: 185-198
Marras R (2002) Analisi di espressione della mutazione di pomodoro parthenocarpic
fruit-2, costituzione di una popolazione segregante F2 e fasi preliminari della
ricerca di marcatori molecolari associati al locus. Graduation thesis, Università
della Tuscia, Viterbo, Italy. Tutor Prof. A Mazzucato
Martiniello P (1975) Effetti biochimici di alcuni geni “Never ripe” sulla bacca di
pomodoro. Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor prof. GP Soressi
Martiniello P and Soressi GP (1977) Valutazione dell’attitudine combinatoria generale e
specifica di linee bs di pomodoro. Annali ISPORT 8, pubbl. 138, pp.12.
Martiniello P, Falavigna A, Soressi GP (1985) Influence of the tomato (Lycopersicon
esculentum Mill.) brown seed (bs) gene on plant and fruit characteristics. Genet.
Agr. 39: 417-422.
Mazzucato A, Pepponi F, Chioccia G, Soressi GP (1998a) Ricostituzione e
valorizzazione della varietà locale di pomodoro Scatolone di Bolsena. Italus Hortus
3: 30-35
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
91
Mazzucato A, Taddei AR, Soressi GP (1998b) The parthenocarpic fruit (pat) mutant of
tomato (Lycopersicon esculentum Mill.) sets seedless fruits and has aberrant anther
and ovule development. Development 125: 107-114
Mazzucato A, Caccia R, Soressi GP (1998c) Expression of the pat syndrome and other
floral traits in a tomato BC1 population after interspecifìc cross with L. pennelli.
Tomato Genet Coop Rep 48: 33-36
Mazzucato A, Testa G, Biancari T, Soressi GP (1999) Effect of gibberellic acid
treatments, environmental conditions, and genetic background on the expression of
the parthenocarpic fruit mutation in tomato. Protoplasma 208: 18-25
Mazzucato A, Olimpieri I, Ciampolini F, Cresti M, Soressi GP (2003) A defective
pollen-pistil interaction contributes to hamper seed set in the parthenocarpic fruit
tomato mutant. Sex Plant Reprod 16: 157-164
Mazzucato A, Mosconi P, Siligato F, Picarella ME, Soressi GP (2006) Caratterizzazione
di varietà locali di pomodoro coltivate nella regione Lazio. Italus Hortus 13: 735740
Mazzucato A, Olimpieri I, Rossi M, Caccia R, Soressi GP (2006) A new reporter
construct to monitor IAA dynamics during tomato development. Tomato Genetics
Cooperative Report 56: 26-28
Mazzucato A, Picarella ME (2007). Tomato bruschetta. SOL Newsletter 17: 10
Mazzucato A, Olimpieri I, Soressi GP (2007) Bruschette a colori? Sì, grazie. Il La 1: 7476
Mazzucato A, Papa R, Bitocchi E, Mosconi P, Nanni L, Negri V, Picarella ME, Siligato
F, Soressi GP, Tiranti B, Veronesi F (2008a) Genetic diversity, structure and
marker-trait associations in a collection of Italian tomato (Solanum lycopersicum
L.) landraces. Theor Appl Genet 116: 657-669
Mazzucato A, Olimpieri I, Siligato F, Picarella ME, Soressi GP (2008b)
Characterization of genes controlling stamen identity and development in a
parthenocarpic tomato mutant indicates a role for the DEFICIENS ortholog in the
control of fruit set. Physiol Plant 132: 526–537
Mendoza de Gyves E (1995) Messa a punto e verifica di metodi di trasformazione in
fagiolo (Phaseolus spp.). PhD Thesis, Università dellaTuscia, Viterbo, Italy. Tutor
Prof GP Soressi
Mensurati F (1991) Selezione di piante di pomodoro insensibili a filtrato colturale di
Phytophtora infestans in varietà parzialmente resistenti al fungo. Graduation thesis,
Università della Tuscia, Viterbo, Italy. Tutor Dott. Paola Crinò
Mensurati F (1994) Induzione di variabilità in pomodoro tramite rigenerazione in vitro,
mutagenesi chimica e loro combinazioni. PhD thesis, Università della Tuscia,
Viterbo, Italy. Tutor Prof. GP Soressi
Minoggio M, Bramati L, Simonetti P, Gardana C, Iemoli L, Santangelo E, Mauri PL,
Spigno P, Soressi GP, Pietta PG (2003) Polyphenol pattern and antioxidant activity
of different tomato lines and cultivars. Ann Nutr Metab 47:64-69
Monti L, Santangelo E, Corrado G, Rao R, Soressi GP, Scarascia Mugnozza GT (2004)
Il “San Marzano”: problematiche e prospettive in relazione alla sua salvaguardia e
alla necessità di interventi genetici. Agroindustria 3: 161-169
Moretti F (1982) Valutazione di tecniche di produzione del seme di cipolla. Graduation
Thesis, Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Motto M, Soressi GP, Salamini F (1975) Mutation frequencies and chimeric formation
in Phaseolus vulgaris after EMS treatment of dormant seeds. Rad Bot 15: 291-29
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
92
Motto M, Soressi GP, Salamini F (1979a) Growth analysis in a reduced leaf mutant of
common bean (Phaseolus vulgaris L.). Euphytica, 28: 593-600
Motto M, Fadda A, Soressi GP, Salamini F (1979b) Lisa P71: varietà nana di fagiolo da
granella a seme bianco resistente al virus del mosaico comune (BCMV). Ann.
ISPORT 8 (123): 1-5
Murakami Y, Tsuyama M, Kobayashi Y, Kodama H, Iba K (2000) Trienoic fatty acids
and plant tolerance of high temperature. Science 287: 476–479
Mustilli AC, Fenzi F, Ciliento R, Alfano F, Bowler C (1999) Phenotype of the tomato
high pigment-2 mutant is caused by a mutation in the tomato omolog of
DEETIOLATED1. Plant Cell 11: 145–157
Odoardi M, Gentinetta E, Soressi GP, Salamini F (1976a) Metabolismo del saccarosio
nel seme di fagiolo a diversi stadi dello sviluppo. Annali ISPORT, Salerno 4: 57-66
Odoardi M, Gentinetta E, Soressi GP, Salamini F (1976b) Forme multiple della
fosfoesoisomerasi nel seme di fagiolo durante lo sviluppo. Annali ISPORT, Salerno
4: 69-71
Odoardi M, Palmieri S, Giacomozzi G, Soressi GP, Salamini F (1976c) Enzimi del
metabolismo dei carboidrati durante lo sviluppo del seme in pisello liscio e rugoso.
Riv. Agron. 10: 233-239
Olimpieri I (2006) Aspetti genetici e molecolari della partenocarpia in pomodoro. PhD
thesis, Università della Tuscia, Viterbo, Italy. Tutor Prof. GP Soressi
Olimpieri I, Siligato F, Caccia R, Mariotti L, Ceccarelli N, Soressi GP, Mazzucato A.
(2007) Tomato fruit set driven by pollination or by the parthenocarpic fruit allele
are mediated by transcriptionally-regulated gibberellin biosynthesis. Planta 226:
877-888
Olimpieri I, Mazzucato A. Phenotypic and genetic characterization of the pistillate
mutation in tomato. Submitted to Theor Appl Genet
Palmieri S, Martiniello P, Soressi GP (1975) Effetti di alcuni geni “Never Ripe” e “High
Pigment” sui principali pigmenti della bacca di pomodoro (L. esculentum Mill.).
Annali Istituto Sperimentale per l’Orticoltura, Salerno, VII: 1-12
Palmieri S, Sozzi A, Gorini FL, Soressi GP (1976) Influenza dei geni Nr e nor sulla
qualità e conservabilità del pomodoro (L. esculentum Mill). Annali I.V.T.P.A., 6:
167-178 (oral comunication presented at Convegno Annuale SIGA, 9-11 October,
1975, Milan-Minoprio)
Palmieri S, Martiniello P, Soressi GP (1978) Chlorophyll and carotene content in high
pigment and green flesh fruits. Tomato Genet Coop Rep 28: 10
Palmieri S, Soressi GP (1978) Effetti di alcuni geni Never ripe e high pigment su
colorazione e valore vitaminico della bacca di pomodoro. I Conv. Naz. “Problemi e
prospettive del pomodoro da mensa”, Monsampolo del Tronto, 7-8/11/1978
I.SP.ORT. Salerno, 95-99
Palmieri S, Odoardi M, Soressi GP, Salamini F (1978) Indoleacetic acid oxidase activity
in to high peroxidase tomato mutants. Physiol Plant 42: 85-90
Paniconi G (2006) Caratterizzazione morfologica e molecolare di una varietà di fagiolo
(Phaseolus vulgaris L.) di origine siciliana. Graduation Thesis, Università della
Tuscia, Viterbo, Italy. Tutor Prof. A. Mazzucato
Pardossi A, Frangi P, Tognoni F, Soressi GP (1988) Growth analysis of monostem
tomato genotype in N.F.T. Acta Hortic 229: 361-369
Parsons EP (2000) Sviluppo di un protocollo di rigenerazione in vitro ai fini della
trasformazione genetica del fagiolo (Phaseolus coccineus L. e Phaseolus vulgaris
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
93
L.). Graduation thesis, Università della Tuscia, Viterbo, Italy. Tutor Prof GP
Soressi
Pecaut P, Philouze J (1978) A sha pat line obtained by natural mutation. Rep Tom Genet
Coop 28: 12
Pepponi F (1995) Ricostituzione e valorizzazzione dell’ecotipo di pomodoro “Scatolone
di Bolsena”. Graduation thesis, Univ. Tuscia, Viterbo. Tutor prof. GP Soressi
Perri E (1991). Effetto della composizione del substrato, del tipo di espianto e del
genotipo sulla frequenza di rigenerazione e di variazione indotta in vitro in
pomodoro. Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor Prof. GP Soressi
Philouze J (1985) Parthénocarpie naturelle chez la tomate. II. Etude d'une collection
variétale. Agronomie 5: 47-54
Philouze J, Pecaut P (1986) Parthénocarpie naturelle chez la tomate. III Etude de la
parthénocarpie due au gène pat (parthenocarpic fruit) de la lignée «Montfavet
191». Agronomie 6: 243-248
Picarella ME, Antonelli M, Astolfi S, Zuchi S, Vernieri P, Soressi GP (2007) Ethylene
and ABA cross-communication and plant growth response to salt stress in tomato
(Solanum lycopersicum L.). Advances in Plant Ethylene Research: Proceedings 7th
International Symposium on the Plant Hormone Ethylene (Pisa, 18-22 June 2006):
113-114
Poma G (1992). Selezione di linee monostelo-partenocarpiche utili ai fini del
miglioramento genetico del pomodoro da tavola e da industria. Graduation thesis,
Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Porcelli S, Soressi GP, Lunicini L (1979) Miglioramento genetico del pomodoro da
mensa. Colture Protette 8: 19-24
Portesi A (1987) Analisi fenotipica e genetica di mutanti della buccia in pomodoro (L.
esculentum Mill.). Graduation thesis, Università Cattolica Sacro Cuore, Piacenza,
Italy. Tutor Prof. GP Soressi
Portesi A, Frangi P, Soressi GP (1987) Allelism test between the fruit skin mutants p,
nor3, pe and pe-2. Tomato Genet Coop Rep 37: 59
Portesi A, Soressi GP (1988) Analisi fenotipica e genetica di mutanti della buccia in
pomodoro. Proceedings of the Symposium on “Physiology of fruit drop, ripening,
storage and post-harvest processing of fruits”, Turin, Italy, pp. 128-135
Quattrini E (1989) Moltiplicazione vegetativa in vitro di cloni diplo-aploidi in asparago.
Graduation thesis, Università Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Ranalli P, Parisi P, Soressi GP, Habashi AA, Santangelo E (2001) Fagiolo (Phaseolus
vulgaris L.) in Leguminose e agricoltura sostenibile: specie da granella e cover
crops. Calderoni Ed agricole, Bologna, pp. 447-502
Ranieri R, Stancanelli G, Tonelli A, Ricci A, Soressi GP (1996) Variabilità somaclonale
in pomodoro da industria (L. esculentum Mill). Italus Hortus 3: 14-21
Remotti P (1990) Variabilità in popolazioni derivate da incrocio Solanum tuberosum L.
x S. Berthaultii per caratteri morfologici e per densità di tricomi ghiandolari
correlati con la resistenza agli insetti. Graduation thesis, Università della Tuscia,
Viterbo, Italy. Tutor Prof. GP Soressi
Ryan CA (1990) Proteinase inhibitors in plants: Genes for improving defenses against
insect and pathogens. Annu Rev Phytopathol 28: 425-449
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
94
Rick CM, Uhlig JW, Jones AD (1994) High alpha-tomatine content in ripe fruit of
Andean Lycopersicon esculentum var. cerasiforme: Developmental and genetic
aspects. Proc Natl Acad Sci USA 91: 12877-12881
Rossi F (1977) Influenza dei geni “Never ripe” sulla conservabilità delle bacche di
pomodoro (L. esculentum Mill.). Graduation thesis, Università Cattolica Sacro
Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Rossi M (2003) Caratterizzazione fenotipica e istologica di mutanti di pomodoro
coinvolti nel metabolismo dell’auxina mediante costrutti reporter auxino-inducibili.
PhD Thesis, Università della Tuscia, Viterbo, Italy. Tutor Prof. GP Soressi
Rotino GL, Schiavi M, Manera M, Roversi N, Casarotti D, Soressi GP (1985) Impiego
di calore di acque residue da centrali termoelettriche nella coltura di cultivar di
fragola neutro-diurna. Colture Protette 14: 51-56
Ruiu F (2006) Integrazione della mutazione positional sterile-2 (ps-2) nella mappa
molecolare di pomodoro. Graduation Thesis, Università della Tuscia, Viterbo,
Italy. Tutor Prof. A Mazzucato
Salamini F, Maggiore T, Lorenzoni C, Soressi GP (1975) Ibridi di pomodoro. III.
Attitudine combinatoria di 86 varietà da industria. Sementi Elette 21: 3-23
Salamini F, Fadda A, Allavena A, Motto M, GP Soressi (1978) Nuove varietà di fagiolo
resistenti al mosaico comune (BCMV) costituiti in Italia. Informatore Agrario 34
(50): 3990-3
Santangelo E (1988) Espressione del gene pat-2 di partenocarpia in pomodoro in
condizioni di fluttuazioni termiche e suo effetto su precocità e stabilità della
produzione. Graduation thesis, Università della Tuscia, Viterbo, Italy. Tutor Prof.
GP Soressi
Santangelo E, Soressi GP (1990) La partenocarpia nel pomodoro. Colture Protette 3: 2933
Santangelo E, Benedettelli S, Tomassini C, Soressi GP (1990) Comportamento in tunnel
freddo delle versioni, quasi isogeniche, normale e partenocarpica (pat-2) di tre
ibridi di pomodoro. Colture Protette 11: 81-90
Santangelo E, Balestra GM, Varvaro L, Soressi GP (1998) Ottenimento di piante di
pomodoro tolleranti sia a Fenthion che a Pseudomonas syringae pv. tomato a
seguito di rigenerazione in vitro, in presenza di Fenthion, di espianti cotiledonari
eterozigoti per il gene Pto di resistenza al batterio. Atti Giornate Fitopatologiche,
Scicli e Ragusa 3-7 Maggio 1998, pp. 719-724
Santangelo E, Balestra GM, Varvaro L, Soressi GP (2000) Efficiency of the in vitro
selection for R4 tomato plants resistant or susceptible to both fenthion and
Pseudomonas syringae pv. tomato. J Plant Pathol 82: 83
Santangelo E, Lai A, Fonzo V, Mosconi P, Fantoni R, Soressi GP (2002) Studio
dell’evoluzione di clorofilla, carotenoidi e polifenoli durante la maturazione della
bacca in linee di pomodoro diversificate per geni del colore, mediante analisi
ottiche. Proceed VI Giornate Scientifiche (vol. II), April 23-25, Spoleto, Italy, pp
471-472
Santangelo E, Fonzo V, Astolfi S, Zuchi S, Caccia R, Mosconi P, Mazzucato A, Soressi
GP (2003a) The Cf-2 / Rcr3esc gene interaction in tomato (Lycopersicon
esculentum) induces autonecrosis and triggers biochemical markers of oxidative
burst at cellular level. Funct Plant Biol 30: 1117-1125
etic
TOMATO GEN omic S [29-30 Maggio 2008,Viterbo – Università della Tuscia]
95
Santangelo E, Habashy AA, Caccia R, Mosconi P, Picarella ME, Mazzucato A, Soressi
GP (2003b) Triploidi partenocarpici di pomodoro: una innovazione nel futuro del
vivaismo orticolo. Italus Hortus 10: 191-194;
Santangelo E, Mazzucato A, Picarella ME, Mosconi P, Lioi L, Soressi GP (2006).
Caratterizzazione del ‘Fagiolo del Purgatorio’ di Gradoli (VT). Italus Hortus 13:
496-502
Santangelo E, Antonelli A, Picarella ME, Soressi GP (2007a) The Lesion Mimic
Mutants as a tool for unveiling the genic network operating during biotic and
abiotic plant stresses. In: Thangadurai D, Tang W, Song SQ (eds) Plant Stress and
Biotechnology (ISBN 978-81-89473-10-5). Oxford Book Company, Jaipur, India,
pp 155-176
Santangelo E, Marabottini R, Antonelli M, Badiani M, Pasqualini S, Soressi GP (2007b)
The autonecrosis triggered by Cf-2/Rcr3esc interaction in tomato (Solanum
lycopersicum L.) involves an ethylene burst and the enzymatic machinery
protecting the respiratory apparatus of mitochondria. In: Ramina A et al. (eds)
Advances in Plant Ethylene Research: Proceedings of 7th International Symposium
on the Plant Hormone Ethylene (Pisa, 18-22 June 2006): 391-392
Schettino M, Caccia R, Testa G, Soressi GP (1998) Piante di pomodoro transgeniche per
l'inibitore di protease Kunitz, finalizzate al controllo dell'Helicoverpa armigera.
Atti IV Giornate Scientifiche SOI, Sanremo, Imperia,1–3 Aprile, pp 505-506
Schettino M (2004) Trasformazione genetica del pomodoro con il gene inibitore di
proteinasi Kunitz (KTI3) e suo effetto su sviluppo e sulla capacità riproduttiva di
Helicoverpa armigera (Lepidoptera noctuidae) Graduation thesis, Università della
Tuscia, Viterbo, Italy. Tutors Proff. GP Soressi and C Pucci
Schiavi M (1977). Eredità di alcuni mutanti che determinano assenza di germogli laterali
nella pianta di pomodoro (L. esculentum Mill.). Graduation thesis, Università
Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Schiavi M, Soressi GP, Maggiore T, Casarotti D, Porcini F, Smedile E (1983)
Utilizzazione del calore residuo da centrali termoelettriche per produzioni vegetali
agrarie: risultati delle esperienze condotte a Tavazzano. Atti 1a Conferenza
Internazionale Energia e Agricoltura, Milano 27-28-29 Aprile 1983 vol. 3-28: 1-25
Schiavi M, Rotino F, Parrini F, Casarotti D, Falavigna A, Maggiore T, Smedile E,
Soressi GP (1984a) Use of waste heat from an electric generating plant (wet tower
cooling) in agricolture: first results with Gerbera cultivation in glasshouse.
Working Group “waste heat utilization” of E.S.N.A. Proceeding of the meeting
held Piacenza 3-7 September 1984: 39-54
Schiavi M, Rotino GL, Parrini F, Casarotti D, Falavigna A, Maggiore T, Smedile E,
Soressi GP (1984b) Use of waste heat from an electric generatine plant (open
circuit cooling) in agricolture: two years of experiments with inbreds of maize.
Proceedings E.S.N.A. working group “waste heat utilization” Piacenza 3-7
September 1984: 55-67
Schiavi M, Casarotti D, Maggiore T, Rotino GL, Soressi GP (1984c) Utilizzazione del
calore delle acque residue della centrale termoelettrica di Tavazzano: un triennio di
prove con fragola. Colture Protette 7: 31-38
Schiavi M, Masera R, Fantino MG, Lorenzoni C, Soressi GP (1984d) Metodi di
selezione per tolleranza a Fusarium e per caratteristiche merceologiche del bulbo in
cipolla. Atti Convegno GOF-SOI, 9 December 1983 Ferrara, pp. 27-30
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Schiavi M, Masera R, Nervo G, Mazzola P, Soressi GP (1984e) Different efficiency of
blow-flies and honey bees in controlled pollination of onion (Allium cepa L.).
Eucarpia 3rd Allium Symposium, Wageningen, The Netherlands 4-6 Sept 1984, p
115
Schiavi M, Masera R, Lorenzoni C, Soressi GP (1988a) Metodi utilizzati per il
miglioramento genetico della Dorata di Parma. Agricoltura e Ricerca 82: 21-30
Schiavi M, Masera R, Nerva G, Mazzola P, Soressi GP (1988b) Diversa efficacia
pronuba dell’ape e della mosca cartaria in condizioni di impollinazione controllata
della cipolla. Agricoltura e Ricerca 82: 35-38
Schiavi M, Vicini E, Tisselli V (1991) Miglioramento genetico: nuove varietà di cipolla.
Agricoltura 19: 64-66 (Regione Emilia Romagna)
Schiavi M, Lorenzoni C, Soressi GP (1998) Onion. In: Italian Contribution to Plant
Genetics and Breeding, GT Scarascia Mugnozza & MA Pagnotta (eds), pp 477480. University of Tuscia, Viterbo
Scolari G (1987). Possibilità e limiti del genotipo “monostelo” nel miglioramento
genetico del pomodoro. Graduation thesis, Università Cattolica Sacro Cuore,
Piacenza, Italy. Tutor Prof. GP Soressi
Sgherri C, Navari-Izzo F, Pardossi A, Soressi GP, Rizzo R (2007) The influence of
diluted sea water and ripening stage on the content of antioxidants in fruits of
different tomato genotypes. J Agric Food Chem 55: 2452-58
Somerville C, Browse J (1991) Plant lipids: metabolism, mutants, and membranes.
Science 252: 80–87
Soressi GP (1964) Effetti genetici dei raggi X su alcune cultivar di pomodoro.
Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. A
Bianchi
Soressi GP (1966) Eterosi nel pomodoro e possibilità di ottenere ibridi senza
demasculazione utilizzando marcatori genetici. Sementi Elette 12: 2-12
Soressi GP, Cravedi P, Marchesi G (1966) Descrizione di mutanti artificialmente indotti
in cultivar di pomodoro. Genetica Agraria 20: 204-219
Soressi GP (1967a) Brown seed (bs) a tomato seed character which behaves as an
endosperm trait. Tomato Genet Coop Rep 17: 50
Soressi GP (1967b) Spongy seed (ss), a new plant character which modifies the seed hair
appearance. Tomato Genet Coop Rep 17: 49-50
Soressi GP, Cravedi P (1967) Tomato mutants obtained by means of X-ray and
ethylmethanesulphonate (EMS) treatments. Tomato Genet Coop Rep 17: 51
Soressi GP (1968) Development of hybrid tomato seed making simultaneous use of the
genetic markers brown seed and spongy seed. Tomato Genet Coop Rep 18: 37-38
Soressi GP (1969a) Il pomodoro. Trattato di genetica agraria speciale. Ed agricole,
Bologna, 47 pp
Soressi GP (1969b) Utilità di alcuni caratteri mutanti nella costituzione di cultivar e di
ibridi per la raccolta meccanica della coltura del pomodoro. Sementi Elette 15: 7681
Soressi GP (1969c) I problemi e le prospettive della meccanizzazione della coltura del
pomodoro con particolare riguardo alle operazioni di raccolta. Atti Gionate della
Meccanizzazione della coltura del pomodoro, Camera di Comm di Piacenza, pp 515
Soressi GP (1970a) Tomato mutants following EMS seed treatments. Tomato Genet
Coop Rep 20: 59
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Soressi GP (1970b) Tomato hybrid seed production: a further scheme. Tomato Genet
Coop Rep 20: 58-59
Soressi GP, Lorenzoni C (1970) Crossing rate after natural and hand pollination with
and without emasculation. Tomato Genet Coop Rep 20: 61-63
Soressi GP, Borghi B, Lorenzoni C, Maggiore T, Mariani BM, Salamini F (1973) Ibridi
di pomodoro determinati. II. Valutazione dell’attitudine combinatoria. Sementi
Elette 19: 3-35
Soressi GP (1974) Pomodoro. In: Aspetti genetici dell’attività sementiera. Monografie
di Genetica Agraria. Bianchi A, Pavia, II: 33-53
Soressi GP, Lorenzoni C, Borghi B (1974a) Valutazione di varietà e ibridi di anguria
(Citrullus vulgaris L. Schrad) quale premessa al miglioramento genetico della
specie. Sementi Elette 20: 9-19
Soressi GP, Gentinetta E, Odoardi M, Salamini F (1974b) Leaf peroxidase activities in
tomato mutants affecting plant morphology. Biochem Genet 12: 181-198
Soressi GP (1975a) New spontaneous or chemically induced fruit ripening tomato
mutants. Tomato Genet Coop Report 25: 21-22
Soressi GP (1975b) Un nuovo ideotipo di pomodoro per la raccolta meccanica. Atti
Giornata orticola sul pomodoro da industria, Grosseto, Settembre 1974. Annali
ISPORT, 6, 1-7
Soressi GP, Salamini F (1975) A monomendelian gene inducing parthenocarpic fruits.
Rep Tom Genet Coop 25: 22
Soressi GP, Palmieri S, Falavigna A (1975) Una nuova tecnica per la produzione di
seme ibrido di pomodoro. Atti della Giornata delle sementi orticole, Ascoli Piceno,
15-18 maggio 1975
Soressi GP (1976) Miglioramento genetico del pomodoro da industria. Conoscere per
produrre 3: 7-12
Soressi GP (1977) Utilizzazione dei marcatori genetici. Riv di Agronomia 11: 52-57
Soressi (1979) Potential of mutation breeding in tomato. Israel-Italian joint meeting on
genetics and breeding of crop-plants. Monografia di Genetica Agraria, IV: 245-258
Soressi GP, Falavigna A (1979) Il miglioramento genetico dell’asparago (Precoce
d’Argenteuil). Atti giornate studio Coltura Asparago, Santena (TO), 13 Maggio
1978, pp. 7-11
Soressi GP, Schiavi M, Tosi A, Marchesi E (1981) Utilizzazione in orticoltura di acque
calde reflue da centrali termoelettriche. Colture Protette 10: 15-19
Soressi GP, Rotino G, Schiavi M, Casarotti D (1985) Utilizzazione delle energie
alternative: le acque reflue tiepide. Italia Agricola 122: 164-171
Soressi GP (1986a) Induced mutations and breeding in vegetable crops. Proc HungarianItalian Plant Genet Conf, June 26-30, pp 17-32
Soressi GP (1986b) Obiettivi e linee di sviluppo delle ricerche sull’asparago.
Agricoltura Ricerca 8: 7-12
Soressi GP, Bruzzone GB, Frangi P (1986) Influenza della densità di coltivazione sulle
caratteristiche morfologiche e produttive di genotipi di pomodoro monostelo e
normale. Annali Facoltà di Agraria, Università Cattolica S. Cuore, Milano 26(1), 2941.
Soressi GP (1988a) Progetto finalizzato MAF Orticoltura: sub-progetto Asparago e
Cipolla. Agricoltura e Ricerca 10(92):43-60
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Soressi GP (1988b) Progetto finalizzato orticoltura: sub progetto asparago e cipolla.
Agricoltura e Ricerca 92: 43-60. Relazione presentata al convegno nazionale
"Progetto finalizzato Orticoltura" Paestum (SA), 10-11 Novembre 1988
Soressi GP, Scarascia Mugnozza GT (1988) Energy saving through new plant
architecture. Acta Hortic 229: 39-50
Soressi GP, Pardossi A, Portesi A (1988) "Monostem" tomato: a plant model for the
high technology Era. International Symposium on High Tecnhology in Protected
Cultivation, Hamamatsu, Japan, May 12-15, 1988 (Book of abstract 2-3)
Soressi GP, Porcelli S (1988) Obiettivi e linee di sviluppo delle ricerche sulla cipolla.
Agricoltura e Ricerca 82: 1-4
Soressi GP, Schiavi M, Falavigna A (1991) Le colture in vitro e la micropropagazione
per il rinnovamento del vivaismo e della ortoflorofrutticoltura. Terra e Vita,
Speciale micropropagazione, 1: 78-82
Soressi GP (1992) Origine, evoluzione, importanza del pomodoro coltivato. pp. 129146. In “Atti Convegno Internazionale "Scambi Floristici fra Vecchio e Nuovo
Mondo. Riflessi Agro-Selvicolturali e impatti naturalistico-ambientali e
paesaggistici". Genova, 22-23 Aprile 1991. Fondazione Reg. Cristoforo Colombo,
Comune - Ente fiera Genova
Soressi GP, Mapelli S (1992). Genetical, physiological and biochemical aspects of a
monostem tomato phenotype suitable for mechanization and processing. Acta
Horticulturae, 301, 229-236
Soressi GP, Campolmi C, Pardossi A, Tognoni F (1992) Nuovi modelli di piante per le
moderne tecniche colturali. Colture Protette 21 (3): 51-57
Soressi GP (2003) Variabilità disponibile e utilizzabile nel miglioramento genetico del
carciofo. L’Informatore Agrario 22: 47-50
Soressi GP, Papalini P, Piccioni C, Santangelo E, Mosconi P, Picarella EM, Mazzucato
A (2007) Innovazione varietale e programmazione colturale. In atti del convegno
“Ricerca ed innovazione per la valorizzazione del carciofo nella regione Lazio”,
ARSIAL Tarquinia 30 novembre 2007: 12-23
Soressi GP, Cammareri G, Picarella MP (2008) Improvement of in vitro vegetative
propagation technique in tomato (Solanum lycopersicum). Acta Horticulturae (in
press)
Sozzi A (1969) Impiego del carattere seme bruno (bs) nella produzione di seme ibrido di
pomodoro. Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor Prof. GP Soressi
Sozzi A, Gorini FL, Soressi GP (1981) Performance of nor2 hybrids of tomato
(Lycopersicon esculentum Mill.) during storage at controlled temperature. Acta
Hort 116: 223-232
Speranza S (2001) Dinamica di popolazione della Nottua gialla del pomodoro
(Helicoverpa armigera (Hüb.) Lepidoptera, Noctuidae) nel Centro Italia mediante
applicazione di Traptest® innescate con due diverse miscele feromoniche.
Informatore Agrario 22/2001: 59-62
Stamegna L (2008) Analisi genetiche in un mutante maschiosterile condizionale di
pomodoro (Solanum lycopersicum L.). Graduation Thesis, Università della Tuscia,
Viterbo, Italy. Tutor Prof. A Mazzucato
Stancanelli G (1989) Analisi di crescita a livello di plantula in asparago. Graduation
thesis, Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
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Stancanelli G, Falavigna A, Soressi GP (1990) Growth analysis of seedlings and spears
in different asparagus genotypes. Acta Horticulturae 271: 503-508
Testa G (1998). Ricerca di geni differenzialmente espressi in mutanti partenocarpici di
pomodoro a confronto con linee quasi isogeniche normali. Specialization thesis,
Università della Tuscia, Viterbo, Italy. Tutor Prof. GP Soressi
Testa G, Caccia R, Tilesi F, Soressi GP, Mazzucato A (2002) Sequencing and
characterization of tomato genes putatively involved in fruit set and early
development. Sex Plant Reprod 14: 269-277
Tiburtini S (1992) Variabilità somaclonale indotta nella cv. Alice di pomodoro a seguito
di rigenerazione in vitro di espianti cotiledonari. Graduation thesis, Università della
Tuscia, Viterbo, Italy. Tutor Prof. GP Soressi
Todisco F (1978) Eredità di alcuni mutanti “sundwarf” e loro utilizzazione nel
miglioramento genetico del pomodoro. Graduation thesis. Università Cattolica
Sacro Cuore, Piacenza, Italy. Tutor Prof. GP Soressi
Venezia A (1991) Raccolta e classificazione botanica di germoplasma di asparago
spontaneo in Sicilia: messa a punto delle tecniche di germinazione e
moltiplicazione in vitro. Graduation thesis, Università Cattolica Sacro Cuore,
Piacenza, Italy. Tutor Prof. GP Soressi
Venezia A, Soressi GP, Falavigna A (1993) Aspetti relativi alla valorizzazione di specie
di asparago spontanee in Italia. Agricoltura Ricerca 141: 41-48
Ziliotti A (1984) Interazione tra mutanti di pomodoro che controllano l’emissione di
germogli laterali (ls e to-2) e la formazione partenocarpica della bacca (pat e pat2). Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof.
GP Soressi
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Memorandum
The graduation theses tutored by Prof. GP Soressi not included in the contributions are
here reported:
Francavilla S (1992) Rigenerazione in vitro e trasferimento di geni marcatori via
Agrobacterium tumefaciens in Solanum spp. per il miglioramento genetico della
melanzana. Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor Prof. GP Soressi
Ferrari F (1996) Le nuove piante da fronda: problematiche inerenti la moltiplicazione
vegetativa con particolare riferimento al Pittosporum tenuifolium cv. silver queen.
Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy. Tutor Prof. GP
Soressi
Parodi A (1992) Miglioramento genetico dell’Arginanthemum frutescens attraverso
mutagenesi. Graduation thesis, Università Cattolica Sacro Cuore, Piacenza, Italy.
Tutor Prof. GP Soressi
Pecchioni N (1980) Analisi della sintesi proteica in condizioni di stress termici in
Gerbera jamesonii hybrida H. Bolus. Graduation thesis, Università Cattolica Sacro
Cuore, Piacenza, Italy. Tutors Proff. GP Soressi and N Marmiroli
Santangelo E (1994) Rivalutazione e valorizzazione del cece attraverso il ricorso a
individuazione di un protocollo di trasformazione in vitro per consentire
l’introduzione di geni utili. PhD thesis, Università della Tuscia, Viterbo, Italy. Tutors
Proff. GP Soressi and E Porceddu
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… and the story continues with… your Notes…………
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