Ph.D Jay W Scott

Luis Mejía

Rudy E Teni

Brenda E García

Ana Cristina Fulladolsa

Luis Méndez

P Maxwell

Amilcar Sánchez

V V Rivera

G A Secor

Christopher Lofty

Julian Smith

Pravda Stoeva-Popova

Tomato resistance to bacterial wilt caused by Ralstonia solanaearum E

The Tomato Genetics Cooperative, initiated in 1951, is a group of researchers who share and interest in tomato genetics, and who have organized informally for the purpose of exchanging information, germplasm, and genetic stocks. The Report of the Tomato Genetics Cooperative is published annually and contains reports of work in progress by members, announcements and updates on linkage maps and materials available. The research reports include work on diverse topics such as new traits or mutants isolated, new cultivars or germplasm developed, interspecific transfer of traits, studies of gene function or control or tissue culture. Relevant work on the Solanaceous species is encouraged as well.

Paid memberships currently stand at approximately 94 (includes those paid in 2009 and beyond) from 16 countries.

Cover: Design by Dolly Cummings. Bacterial wilt incited by Ralstonia solanacearum is a serious threat to tomato production in many humid tropical production regions. Breeding for resistance has been a challenge due to multiple strains of the bacterium, variable environmental effects on disease expression, largely unknown genetics for resistance, and linkage of resistance to small fruit size. Locating molecular markers tightly linked to resistance genes should be a boon to future breeding efforts and with the tomato genome now sequenced this research may advance much more rapidly than in the past. This years" feature article explores the major sources ("roots") of bacterial wilt resistance and thereby sheds light on the relationships of genotypes that could be used in studies to locate molecular markers. Precision in the exact identification of sources is hampered by a lack of information or by conflicting information. Unfortunately some of the old sources are no longer available. If you read the article and have information that would help us improve the article please contact us. We live in an electronic age and if we get better information we will put it in the article and future versions may be better than the one published in December of 2010. If you have seed of a rare source line send it to us and we will see that it gets in a gene bank. Thanks for your help.

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Author Index Introduction Bacterial wilt is caused by the pathogen formerly known as Pseudomonas solanacearum, transitorily renamed Burkholderia solanacearum [START_REF] Yabuuchi | Proposal of Burkholderia gen. nov. and transfer of seven species of the genus Pseudomonas homology group II to the new genus, with the type species Burkholderia cepacia (Palleroni and Holmes 1981) comb[END_REF] and presently accepted as Ralstonia solanacearum [START_REF] Yabuuchi | Transfer of two Burkholderia and an Alcaligenes species to Ralstonia gen. Nov.: Proposal of Ralstonia pickettii (Ralston, Palleroni and Doudoroff 1973) comb. Nov., Ralstonia solanacearum (Smith 1896) comb. Nov. and Ralstonia eutropha (Davis 1969) comb[END_REF][START_REF] Vaneechoutte | Wautersia gen. nov., a novel genus accommodating the phylogenetic lineage including Ralstonia eutropha and related species, and proposal of Ralstonia [Pseudomonas] syzygii (Roberts et al. 1990) comb[END_REF]. Developing varieties with resistance has challenged tomato breeders for over 60 years for several reasons. Strong interactions are observed between resistance, environmental conditions and strains (e.g. [START_REF] Kelman | The bacterial wilt caused by Pseudomonas solanacearum. A literature review and bibliography[END_REF][START_REF] Acosta | Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill[END_REF][START_REF] Krausz | Breakdown of resistance to Pseudomonas solanacearum in tomato[END_REF][START_REF] Messiaen | Environmental influences on the severity of tomato bacterial wilt in the French West Indies: Interactions with varietal resistance[END_REF][START_REF] Peter | Breeding for resistance to bacterial wilt in tomato, eggplant and pepper[END_REF][START_REF] Prior | Resistance to bacterial wilt (Pseudomonas solanacearum) in tomato: present status and prospects. In: Bacterial Wilt, the disease and its causative agent Pseudomonas solanacearum[END_REF][START_REF] Hanson | Variable reactions of tomato lines to bacterial wilt evaluated at several locations in Southeast Asia[END_REF][START_REF] Jaunet | Variation in genotype and aggressiveness of Ralstonia solanacearum race 1 isolated from tomato in Taiwan[END_REF][START_REF] Balatero | QTL mapping for bacterial wilt resistance in Hawaii 7996 using AFLP, RGA and SSR markers[END_REF][START_REF] Hai | Resistance to Taiwanese race 1 strains of Ralstonia solanacearum in wild tomato germplasm[END_REF]. Further, several defaults are often associated to bacterial wilt resistance, such as small fruit size [START_REF] Acosta | Heritability of bacterial wilt resistance in tomato[END_REF][START_REF] Gilbert | Horticultural refinement of multiple disease resistant tomatoes in Hawaii[END_REF][START_REF] Scott | Breeding tomatoes for resistance to bacterial wilt, a global view[END_REF], bitterness due to high tomatine content [START_REF] Borchers | Quantitative estimation of a bitter principle in tomato[END_REF][START_REF] Mohanakumaran | Bacterial wilt resistant tomato lines with unusually high content of the alkaloid tomatin[END_REF][START_REF] Digat | A study of the varietal resistance to bacterial wilt[END_REF][START_REF] Messiaen | Cumulate resistances to Pseudomonas solanacearum and to Meloidogyne icognita with determinate growth in tomato[END_REF], green gel around the seeds and epidermis cracking [START_REF] Acosta | Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill[END_REF][START_REF] Cordeil | Etude de la résistance variétale de la tomate au flétrissement bactérien en Guadeloupe et en Guyane française[END_REF][START_REF] Daly | IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies[END_REF]. The difficulty in developing resistance with adaptation to heat and with good horticultural features, in particular fruit firmness and large size, has been reported by many authors (e.g. [START_REF] Kaan | Recherche de variétés de tomates aux Antilles[END_REF] available information on the phenotype of the main resistant accessions lines displayed in Figure 1 and issued from the main historical breeding programs.

Puerto Rico

Information about tomato breeding for bacterial wilt resistance in the University of Puerto Rico is very scarce and dispersed in annual reports of the Puerto Rico University Agricultural Experiment Station. For instance, [START_REF] Cook | Annual report of the division of botany and plant pathology (tomatoes)[END_REF][START_REF] Cook | Annual report of the division of botany and plant pathology (wilt disease of tomato)[END_REF], [START_REF] Roque | Annual Report of the Assistant Pathologist (wilt disease of tomatoes)[END_REF] and [START_REF] Theis | Tomato diseases[END_REF] mention partial resistance of some native material and its use in breeding. [START_REF] Warmke | Tomato breeding. Puerto Rico Fed[END_REF] experimented with local selections from hybrids between native and imported tomato varieties, some of which showing some resistance to bacterial wilt and out-yielding the controls. [START_REF] Azzam | Tomato breeding for the tropics[END_REF] reported the existence of resistance in S. pimpinellifolium as well as the development of breeding lines with some degree of resistance but with unacceptable fruit quality.

At the beginning of the 1960s, at Rio Piedras station, a double cross involving ["Platillo" , ( a native variety) X a S. pimpinellifolium, (of unknown identity)] X [a tomato, (of unknown identity) X "Platillo"] was made by H. Azzam [START_REF] Digat | A study of the varietal resistance to bacterial wilt[END_REF]; [START_REF] Daly | IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies[END_REF] and its progenies were used by IRAT in the 1960s and 1970s [see below]. The literature also mentions "Beltville 3814 (=T414)" which was said to be from Puerto Rico and was used in North Carolina breeding effort [See below].

North Carolina

The search and breeding of bacterial wilt resistant tomatoes began long ago in the USA, at the turn of the 19 th and 20 th century in the agricultural experiment stations of several States including North Carolina, Florida, Alabama, and Mississipi [START_REF] Kelman | The bacterial wilt caused by Pseudomonas solanacearum. A literature review and bibliography[END_REF]. After a lapse of years, efforts were resumed at the North Carolina Experiment Station in 1936 [START_REF] Schmidt | Tomato breeding[END_REF][START_REF] Schmidt | Tomato breeding[END_REF] and involved many horticultural scientists such as W.S. Barham, F.D. Cochran, M.E. Gardner, W.R. Henderson, J.S. Weaver, and plant pathologists such as D.E. Ellis, S.F. Jenkins, A. Kelman and N.N. Winstead (Henderson & Jenkins, 1972b). [START_REF] Warmke | Tomato breeding. Puerto Rico Fed[END_REF] as well as [START_REF] Walter | Hereditary resistance to disease in tomato[END_REF] mention the existence of a US Southern Tomato Exchange Program (STEP) that was put into operation in 1946 [START_REF] Yarnell | The southern tomato exchange program[END_REF], and was complemented with the National Screening Program for evaluation of PI (Plant Introduction) accessions of Lycopersicon for disease resistance. These programs are probably at the origin of the complex relationships between the breeding research carried out in North Carolina, Hawaii, Florida and Puerto Rico for bacterial wilt resistance.

The two widely mentioned sources of resistance of North Carolina breeding material are S. lycopersicum var. cerasiforme "PI 129080" (=T 702) from Colombia (initially classified as L. pimpinellifolium - Henderson & Jenkins, 1972b) and (ii) S. lycopersicum var. pyriforme "Beltsville No. 3814" (=T 414) (Henderson & Jenkins, 1972a & b;Laterrot et al., 1978;[START_REF] Hanson | Diallel analysis of bacterial wilt resistance in tomato derived from different sources[END_REF]. "Beltsville No. 3814" also named "P.I. No. 3814" by [START_REF] Kelman | The bacterial wilt caused by Pseudomonas solanacearum. A literature review and bibliography[END_REF] originated in Puerto Rico according to this author and others [START_REF] Winstead | Inoculation techniques for evaluating resistance to Pseudomonas solanacearum[END_REF]Henderson & Jenkins, 1972a & b) without further details. [START_REF] Thurston | Proc. 1rst Int. Conference and Worskshop on the ecology and control of bacterial wilt caused by Pseudomonas solanacearum[END_REF] also said that it is a selection from Puerto Rico. However the name of this line and the fact that it has a PI number both suggest that "Beltsville No. 3814" was associated with the USDA, Beltsville (Maryland). Perhaps USDA researchers were collaborating with Puerto Rican researchers. "T414" displayed, as other bacterial wilt resistant lines, a specific bitter taste with lasting burning sensation, and it was used by [START_REF] Borchers | Quantitative estimation of a bitter principle in tomato[END_REF] for setting up a quantitative chemical test of the alkaloids responsible of this taste. Another source of resistance, S. lycopersicum "Mulua" from Guatemala, is mentioned by [START_REF] Winstead | Inoculation techniques for evaluating resistance to Pseudomonas solanacearum[END_REF] and by [START_REF] Suzuki | Studies on breeding eggplants and tomatoes for resistance to bacterial wilt. I. Investigations on method of evaluating the resistance and on the source of resistance in eggplants and tomatoes[END_REF] [who refer to [START_REF] Winstead | Inoculation techniques for evaluating resistance to Pseudomonas solanacearum[END_REF] as well as to a personal communication of D.E. Ellis]. According to [START_REF] Winstead | Inoculation techniques for evaluating resistance to Pseudomonas solanacearum[END_REF], "Mulua" yielded resistant breeding material after an initial cross with "Rutgers". Suzuki et al. (1964, p.99) still referring to [START_REF] Winstead | Inoculation techniques for evaluating resistance to Pseudomonas solanacearum[END_REF], mention also a "T-141" from Puerto Rico as another source of resistance used in North Carolina, though these latter authors mention "T414" and not "T-141". The identity of "T-141" is henceforth doubtful, either a mistyping of "T414" by [START_REF] Suzuki | Studies on breeding eggplants and tomatoes for resistance to bacterial wilt. I. Investigations on method of evaluating the resistance and on the source of resistance in eggplants and tomatoes[END_REF] or another line not reported in any other source we found.

To sum up, the cherry tomato "PI 129080" (=T 702) from Colombia, the pear shaped tomato "Beltsville No. 3814" (=T 414) and the tomato "Mulua" from Guatemala, have been included in NC breeding programs as genitors of bacterial wilt resistance.

"Venus" and "Saturn", released in the early 1970s (Henderson & Jenkins, 1972a & b) are the best known commercial varieties with resistance to bacterial wilt issued from the North Carolina State program. Their pedigrees are provided in Fig. 2, and their phylogenic relationship with two other NC lines, "MR4" and "NC 72 TR 4-4", is provided in Fig. 3. Hawaii D.C. McGuire, J.C. Gilbert and J.C. Acosta (breeders) as well as I.W. Buddenhagen (pathologist) are among the scientists having worked on tomato resistance to bacterial wilt in the course of the 1950s and 1960s. Breeding for resistance in commercial type tomatoes was confined first (prior to 1955) to crossing root knot nematode resistant Hawaii lines and North Carolina bacterial wilt tolerant lines [START_REF] Acosta | Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill[END_REF][START_REF] Acosta | Heritability of bacterial wilt resistance in tomato[END_REF]. [START_REF] Acosta | Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill[END_REF] indicated that several North Carolina lines which had been bred for bacterial wilt resistance, proved to be intermediate in wilt susceptibility under Hawaiian conditions. In 1953 a new source of resistance, S. pimpinellifolium "PI 127805A", originating from Peru, was "obtained" [sic]5 and field selected in Hawaii through 9 generations [START_REF] Acosta | Heritability of bacterial wilt resistance in tomato[END_REF]. This accession would be at the origin of the line "5808-2" [START_REF] Mohanakumaran | Bacterial wilt resistant tomato lines with unusually high content of the alkaloid tomatin[END_REF]. [START_REF] Acosta | Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill[END_REF] writing the name as "HES 5808-2", mentions that it is an inbred line of L. pimpinellifolium obtained by D.C. McGuire, but he does not refer to any PI number. The commercial variety "Kewalo" (Fig. 4), developed by [START_REF] Gilbert | Kewalo" tomato[END_REF], recombines the resistance to bacterial wilt originating from "PI 127805A" with root-knot nematode resistance(gene Mi) and other useful traits from "Anahu", a local tomato, and its derivative "Kalohi".

"Hawaii 7996" (H7996), "Hawaii 7997" (H7997) and "Hawaii 7998" (H7998) were later bred by J.C. Gilbert in the 1970s [START_REF] Scott | Breeding tomatoes for resistance to bacterial wilt, a global view[END_REF] and it has been reported in the literature [START_REF] Hanson | Diallel analysis of bacterial wilt resistance in tomato derived from different sources[END_REF][START_REF] Balatero | Identification of AFLP and RGA markers associated with bacterial wilt resistance QTL derived from tomato Lycopersicon pimpinellifolium[END_REF][START_REF] Scott | Breeding tomatoes for resistance to bacterial wilt, a global view[END_REF]) that these lines had resistance derived from PI 127805A. However, correspondence dated October 1983 from J. Tanaka (Assistant Horticulturist at Hawaii University) to J. Scott indicates that the 3 mentioned Hawaiian lines and five others ("H7975", "H7976", "H7981", "H7982", "H7983") are sister lines selected from a initially highly variable accession named "HSBW", which acronym might mean "Hot Set Bacterial Wilt". This enigmatic accession, delivered to J.C. Gilbert via an unidentified way and planted first in 1973, displayed a high level of bacterial wilt resistance in hot tropical areas. An earlier correspondence, dated March 1978, from J.C. Gilbert to H. Laterrot (INRA) indicates that "Hawaii 7996" could be cited as having been selected at the University of Hawaii for bacterial wilt resistance, and as having its origin somewhere in the Philippines. J.C. Gilbert admitted in this letter that no publications had been specifically written for this line, and that he was not fully satisfied with it because of its flavour due to alkaloid residues in the ripe fruit. He recommended to using it as a rootstock or as a parent to be crossed with another parent of good flavour and some bacterial wilt resistance, for making F 1 s. Laterrot et al. (1978) also mentioned as a personal communication of Gilbert, a Philippine source at the origin of "Hawaii 7996". H. Laterrot recorded in his handwritten notes, based on Kaan personal communication, that "Hawaii 7996" was a selection made in "Hotset X Philippine tomato" and that it had very small fruits and a determinate growth habit. Given the absence of further written details, it is probably impossible to unravel further the pedigrees of "HSBW" and "Hawaii 7996".

In conclusion, a close look at the dispersed information relative to the breeding programs of Hawaii University indicates that several sources of resistance have been used successively there, first North Carolina material, then "PI 127805A" and lastly Philippines material. It is possible (or probable), that J.C. Gilbert recombined these sources along the time in his breeding material, in one way or another. Breeding is an art as much as a science, and the exact pedigree of the most famous bacterial wilt resistant line "Hawaii 7996" will probably remain the secret of the breeding genius of the late J.C. Gilbert.

Florida

According to [START_REF] Sonoda | Bacterial wilt of tomato in Florida: history, status, and sources of resistance[END_REF] the first attempts to search for bacterial wilt resistance in Florida started over one century ago, but the breeding efforts took a real momentum in the late 1970s. The original sources of resistance used were "Hawaii 7997", S. lycopersicum var. cerasiforme "CRA 66" and S. lycopersicum "PI 126408". The latter is one of the 28 PI accessions determined as resistant to bacterial wilt out of 909 accessions tested [START_REF] Barham | Sources of resistance to late blight and bacterial wilt[END_REF]. However most of the material derived in Florida utilized "Hawaii 7997". Attempts to pyramid resistance genes in the early 1980"s were not successful as there were no molecular markers to identify the genes in resistant plants. No lines were developed that had resistance greater than any of the sources, so there was no evidence that improvements were made under Florida conditions. Over the years it was evident that it was difficult to attain large fruit with high resistance levels. In 1995 "Neptune" (Fig. 5), a line with larger fruit size than "Hawaii 7997", was released, but its resistance level was less than that of "Hawaii 7997" (Scott et al., 1995a), and when tested in the world wide test (as Fla. 7421) "Neptune" also displayed a much narrower spectrum of resistance [START_REF] Wang | Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato[END_REF]. Breeding efforts then focused on taking lines like "Neptune" and crossing them back to "Hawaii 7997" to attain large fruited lines with high resistance levels. At first, new lines were developed with moderate fruit size and high resistance. The next crosses were with very large fruited susceptible inbreds. From this work the sister lines "Fla. 8109" and "Fla. 8109B" were developed [START_REF] Scott | Development of a large fruited tomato with a high level of resistance to bacterial wilt (Ralstonia solanacearum)[END_REF] and further crossing provided new inbreds with high resistance and very large fruit [START_REF] Scott | High level of resistance to bacterial wilt (Ralstonia solanacearum) obtained in large-fruited tomato breeding lines derived from Hawaii 7997[END_REF]. Hypothetically, "Fla. 8109" and the lines developed thereafter contain a resistance gene, missing in "Neptune", that was unlinked from a gene preventing large fruit formation, but this has not been elucidated yet.

Japan

In Japan, development of bacterial wilt resistance in tomato (and eggplant) started as early as 1951 and was based on the use of North Carolina lines. "OTB-1" and "OTB-2" are self pollinated offspring obtained in the 1950s respectively from "NC1953-60N" and "NC1953-64N" [START_REF] Suzuki | Studies on breeding eggplants and tomatoes for resistance to bacterial wilt. I. Investigations on method of evaluating the resistance and on the source of resistance in eggplants and tomatoes[END_REF]. According to later reports and papers published [in Japanese] by the Ministry of Agriculture and Forestry (H. Fukuoka6 , pers. com.) "OTB-2" was segregating for several traits including bacterial wilt resistance (but was fixed for Fusarium wilt resistance) and was submitted to further screening for bacterial wilt resistance and further selfing. In 1969 "BF-Okitsu 101" was obtained7 from this process. "OTB1" and "OTB2" were described in 1964 by IRAT (French West Indies) as possessing the traits of the Puerto Rican S. pimpinellifolium, but with markedly bigger fruits and exceptional fruit productivity, together with a good behaviour towards viruses.

Philippines

In Philippines, breeding was established as early as 1954 by T.L. York and J.R. Deanon who evaluated local and foreign accessions with known resistance to bacterial wilt [START_REF] Deanon | Biochemical bases of screening tomato for bacterial wilt resistance[END_REF]. The exact origin of the resistance source(s) used in the Philippines breeding scheme is not found in the literature and hence remains confusing [START_REF] Deanon | Biochemical bases of screening tomato for bacterial wilt resistance[END_REF][START_REF] Wang | Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato[END_REF]. [START_REF] Empig | Screening tomato, eggplant, and pepper varieties and strains for bacterial wilt (P. solanacearum E.F.S.) resistance[END_REF] report some resistance in Philippine native material, such as "Los Baños native" which has possibly been used in the local breeding research. J. Acosta8 conducted research on the inheritance of tomato bacterial wilt resistance at the University of Hawaii [START_REF] Acosta | Heritability of bacterial wilt resistance in tomato[END_REF]) and one can hypothesize (i) that he took Philippine material to Hawaii 9 and conversely (ii) that he brought material back home (to the Philippines). North Carolina material entered Philippines breeding program according to [START_REF] Mew | Effect of soil temperature on resistance of tomato cultivars to bacterial wilt[END_REF]. These authors, on the basis of a personal communication of J.R. Deanon, indicate that "Venus" (North Carolina line) entered the pedigree of the Philippine line "UPCA1169", together with a "CA64-1169" of unmentioned origin (Figure 1). "UPCA1169" is itself at the origin of other Philippines material such as "VC8-1-2" and "VC9-1" [START_REF] Mew | Effect of soil temperature on resistance of tomato cultivars to bacterial wilt[END_REF][START_REF] Wang | Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato[END_REF][START_REF] Scott | Breeding tomatoes for resistance to bacterial wilt, a global view[END_REF]. The origin(s) of the resistance of other valuable Philippine material, such as "TML46", "TML114", "R3034", or "HSBW" mentioned in the Hawaii section is not known.

French West Indies (INRA & IRAT) and Burkina Faso (IRAT)

In the French West Indies, the research on tomato bacterial wilt resistance started in 1963 at IRAT, and in 1964 at INRA, with some collaboration between the two institutes.

According to [START_REF] Cordeil | Etude de la résistance variétale de la tomate au flétrissement bactérien en Guadeloupe et en Guyane française[END_REF] a collection from Rio Pedras station of the University of Puerto Rico was introduced at IRAT Guadeloupe beginning of 1964. These authors mention a variable tolerance to bacterial wilt of some lines such as "199 PR", "Platillo 78" and L. pimpinellifolium under local conditions. "199 UPR", a derivative from the double cross made at Puerto Rico University ["Platillo" X S. pimpinellifolium] X [a tomato of unknown identity X "Platillo"] -see above-, was chosen for its good tolerance to bacterial wilt [START_REF] Daly | IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies[END_REF]. After pedigree selection, the F 8 lines "199 UPR -39.15" and "199 UPR -39.16" were obtained by [START_REF] Daly | IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies[END_REF]. These lines were described as having small watery and not fleshy fruits, with greenish gel around the seeds, and displaying many concentric cracks [START_REF] Irat | Annual Report[END_REF][START_REF] Cordeil | Etude de la résistance variétale de la tomate au flétrissement bactérien en Guadeloupe et en Guyane française[END_REF]. Both were crossed with "Floralou", a variety of good quantitative and qualitative yield in the conditions of the French West Indies [START_REF] Cordeil | Etude de la résistance variétale de la tomate au flétrissement bactérien en Guadeloupe et en Guyane française[END_REF][START_REF] Digat | A study of the varietal resistance to bacterial wilt[END_REF][START_REF] Daly | IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies[END_REF]. After pedigree selection, the lines "IRAT L3" [START_REF] Daly | IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies[END_REF]Laterrot et al., 1978) -Fig. 6-and "Farako-Ba" (D"Arondel de Haye, unpubl., [START_REF] Irat | Annual Report[END_REF][START_REF] Irat | Annual Report[END_REF]Laterrot et al., 1978;[START_REF] Rouamba | A case of relation between resistances to Pseudomonas solanacearum and Verticillum pathotype 2[END_REF] were respectively obtained in Martinique and Burkina Faso.

The INRA program in Guadeloupe was based on the use of "CRA66". The origin of this line is still controversial. It is given by [START_REF] Digat | A study of the varietal resistance to bacterial wilt[END_REF] and Anaïs (pers. com.) as one of the many small fruited tomato ecotypes grown in Guadeloupe at that time, and known there as "tomadoses". Digat & Derieux described "CRA66" as a vigorous line bearing small, pink, and bitter fruits with resistance to bacterial wilt. However another origin of CRA66 is suggested by Kaan (pers. com.) as being "OTB2", because the phenotype of "CRA66" is very different from the phenotype of the tomadoses: the plant is more vigorous, the leaves have a spreading leaf growth habit, the flowers display an exerted style, the fruits are fasciated and although of pink colour, they have a larger size than tomadoses fruits. There is a green gel inside the fruit, the taste of which is more acrid and extremely bitter, and its bacterial wilt resistance level is higher. However, [START_REF] Suzuki | Studies on breeding eggplants and tomatoes for resistance to bacterial wilt. I. Investigations on method of evaluating the resistance and on the source of resistance in eggplants and tomatoes[END_REF] reported "OTB1" was pink fruited, whereas they described "OTB2" as red fruited (but segregating for several traits). As IRAT experimented in Guadeloupe "OTB1", "OTB2" and many other bacterial wilt resistant lines in the mid-1960s [START_REF] Irat | Annual Report[END_REF][START_REF] Irat | Annual Report[END_REF], it is plausible that valuable material "reached" INRA by some path and perhaps under a distorted identity for some reason. If "CRA66" = "OTB2", then all the French West Indies material would derive indirectly from North Carolina material -see Fig. 1-. Comparison of molecular fingerprinting of "OTB2" and "CRA66" is necessary for elucidating the identity of these two lines.

Crosses started in the 1960s between "CRA66" and the susceptible commercial type "Floradel" resulted in the varieties "Cranita" (Fig. 7), "CRA74" & "Carette" (Fig. 8 &9), and "Caraibo" (= "Caraibe") -Fig. 9-that were respectively released in 1971, 1973[START_REF] Irat | Annual Report[END_REF][START_REF] Valkova-Achkova | L. peruvianum a source of CMS[END_REF][START_REF] Anaïs | Utilisation de la résistance variétale dans la lutte contre le flétrissement bactérien de la tomate Pseudomonas solanacearum E.F. Smith[END_REF][START_REF] Anaïs | La tomate. Pp[END_REF]. "CRA84-26-3" and "Caravel" are offspring of the cross ["Caraibo" X "HC8"] where "HC8" is a heat-tolerant and bacterial wilt resistant line derived from the cross "Hawaii 7996" x "Campbell 28". Later on "Caraibo", "HC8" and "Caravel" were used as genitors in a recurrent selection for recombining their bacterial wilt resistance and agro-climatic adaptation to French West Indies conditions, together with resistance to Begomoviruses [START_REF] Ano | Création de variétés de tomate résistantes à Ralstonia solanacearum et PYMV, adaptées aux régions tropicales[END_REF][START_REF] Ano | L"amélioration de la tomate pour les régions tropicales de plaine: travaux en Guadeloupe. Les familles CRAPY associent les résistances aux Bégomovirus PYMV et TYLCV à la résistance au Ralstonia solanacearum (race 1)[END_REF]. Taiwan AVRDC started tomato breeding in 1972, and from 1973-1980 emphasized the development of breeding lines with heat-tolerance and bacterial wilt resistance [START_REF] Opena | Genetic improvement of tomato adaptability to the tropics: progress and future prospects[END_REF]. Sources of BW resistance frequently used as parents in AVRDC breeding included varieties "Venus" and "Saturn" from North Carolina State University and lines "VC 11-3-1-8", "VC 8-1-2-7", "VC 48-1" from the University of the Philippines. Most AVRDC bacterial wilt resistant lines developed in the 1970"s such as "CL8d-0-7-1" (derived from "VC11-1-2-1B" x "Venus"), "CL9-0-0-1-3" (derived from "VC11-1-2-1B" x "Saturn"), and "CL11d-0-2-1" (derived from "VC9-1-2-9B" x "Venus") were bred from crosses of these two sources. Many AVRDC lines developed in the late 1970"s and early 1980"s such as "CL1131", "CL5915" (Fig. 10), and "CLN65" (Fig. 11) arose from complex crosses involving North Carolina, Philippines, or AVRDC lines with BW resistance derived from the above sources. High levels of BW resistance were detected in "L285", a small-fruited S. lycopersicum germplasm accession from Taiwan (Chang #1) but this source was not used at AVRDC in breeding because it was thought that its bacterial wilt resistance and small fruit size were closely associated [START_REF] Opena | Breeding for bacterial wilt resistance in tropical tomato[END_REF]. In 1985 AVRDC received resistant lines developed in Guadeloupe, including "CRA84-58-1" and "CRA84-26-3", that combined BW resistance and large fruit size. CRA lines were crossed to heat tolerant and BW resistant AVRDC lines which led to the development of AVRDC lines "CLN1462", "CLN1463" (Fig. 12), "CLN1621", "CLN2026", and many others.

Limit of the reliability of the survey: insufficient accuracy of the information

The information found in the literature is often vague and sometimes inconsistent between publications. The absence of published pedigrees for many of the important varieties resistant to bacterial wilt is a real impediment for ascertaining the original sources of their resistance. Henceforth, Figure 1 is the result of our interpretation of sometimes blurred information as exemplified below.

Names of the accessions Depending on the publications, the accessions used in the various trials or breeding programs are not exactly named the same way. This is due in some cases to probable renaming such as for "199 PR" [START_REF] Cordeil | Etude de la résistance variétale de la tomate au flétrissement bactérien en Guadeloupe et en Guyane française[END_REF], which is also found as "199" [START_REF] Digat | A study of the varietal resistance to bacterial wilt[END_REF], "199 UPR" [START_REF] Daly | IRAT L3" une nouvelle variété de tomate combinant plusieurs résistance aux maladies[END_REF], and "UPR 199" [START_REF] Kaan | Recherche de variétés de tomates aux Antilles[END_REF]. The same situation is encountered for "P.I. No. 3814" [START_REF] Kelman | The bacterial wilt caused by Pseudomonas solanacearum. A literature review and bibliography[END_REF], also found as "Beltsville #3814" (Henderson & Jenkins, 1972a;[START_REF] Hanson | Diallel analysis of bacterial wilt resistance in tomato derived from different sources[END_REF], "Beltsville No. 3814" (Henderson & Jenkins, 1972b), and "Beltsville 3814" (Laterrot et al., 1978). However other variations of names such as for "H7997" also found as "H7997S" and "H7997L", "H7998" also found as "H7998S" and "H7998M", or "CRA66" found as "CRA66P" and "CRA66S" in [START_REF] Wang | Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato[END_REF] and [START_REF] Scott | Breeding tomatoes for resistance to bacterial wilt, a global view[END_REF] do not mean further selections but encode only the name of the person who provided the seeds used in the trials. These latter authors mention "BF-Okitsu", otherwise found as "BF-Okitsu 101" in Laterrot (1999). Despite the word "Okitsu" (location in Japan where breeding was carried out, H. Fukuoka, pers. com.) is present in the names of bacterial wilt resistant "BF-Okitsu 101" and of bacterial canker resistant "Okitsu Sozai n°1 10 (Kuriyama & Kuniyasu, 1974), these accessions should not be confused with each other.

Contradictory data found in the literature Apart from the controversy concerning the identity of "CRA66", other contradictory information is found in the literature. Particularly in the case of "Hawaii 7996"and the other Hawaii 79## accessions, resistance was said to originate from a Philippines accession according to several personal communications between breeders in the 1970s, but according to [START_REF] Acosta | Heritability of bacterial wilt resistance in tomato[END_REF] "PI 127805A" and North Carolina material are at the origin of the resistance to bacterial wilt of Hawaiian material. It is not possible to reconcile these conflicting statements other than to say that the various sources of resistance were introduced to the Hawaiian program over time. Therefore we left all options on Figure 1.

Unclear origin of some accessions We found no original information on the origin or pedigree of "Beltsville 3814" and further search of 1960s and 1970s publications of Beltsville USDA research station is needed. Another case concerns the unclear relationship between the Peruvian S. pimpinellifolium "PI 127805" collected in 1938 and maintained by the USDA Northeast Regional PI Station and "PI 127805A" obtained in 1953 and field selected for resistance through 9 generations in Hawaii according to [START_REF] Acosta | Heritability of bacterial wilt resistance in tomato[END_REF]. Furthermore, "5808-2" was derived from "PI 127805A" according to [START_REF] Mohanakumaran | Bacterial wilt resistant tomato lines with unusually high content of the alkaloid tomatin[END_REF] but is given as an inbred line of an anonymous L. pimpinellifolium according to [START_REF] Acosta | Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill[END_REF].

Likeness between accessions Phenotypically "BF-Okitsu" is very close to "Hawaii 7998" (J. Scott and J. Wang, pers. obs.), though the published information (Fig. 1) does not indicate a closer relationship than the presence of North Carolina material in both their pedigrees. Comparison of their molecular fingerprinting would be worthwhile to clarify their genetic relationship.

Sources of resistance and inheritance patterns

The global survey of the major breeding research for tomato bacterial wilt resistance points out that the main sources of resistance used worldwide are perhaps only half a dozen accessions of S. pimpinellifolium ("PI 127805A"), S. lycopersicum var. cerasiforme ("PI 129080"), S. lycopersicum var. pyriforme ("Beltsville 3814"), a progeny from a cross between a S. pimpinellifolium and S. lycopersicum ("199 UPR"), S. lycopersicum ("Mulua") and the enigmatic Philippine accession used in Hawaii in the 1970s. This number could be extended to seven accessions if one adds "CRA66" by assuming it is a Guadeloupe tomadose and not a progeny from "OTB2". Whether there are 6 or 7 main sources of resistance, the genetic basis of tomato resistance mechanisms used worldwide for breeding is quite narrow. North Carolina material has been integrated in most of the other breeding programs, in particular those of Hawaii, Japan, Philippines and Taiwan. By combining NC material, or not, with other sources of 10 "Okitsu Sozai n°1" resistance to Clavibacter michiganensis (formerly named Corynebacterium michiganense) originates from S. habrochaites (L. hirsutum var. glabratum) "PI 134418". resistance, and breeding in geographical areas where different strains of bacterial wilt are prevalent, the breeders exploited the genetic potentialities at their disposal, and created material resisting a wide range of bacterial wilt strains as exemplified by the results obtained in the worldwide trial carried out by [START_REF] Wang | Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato[END_REF]. Indeed, the top nine resistant accessions which had high levels of resistance in almost all 12 locations tested (>90% survival on average) were developed in Hawaii ("H7996", "H7997 S and L", "H7998 S and M"), Philippines ("TML46" and "TML114", "R3034"), and Japan ("BF-Okitsu").

Other sources of resistance in wild tomatoes have been described sporadically in the literature in accessions of the same species (S. pimpinellifolium, cherry and pear S. lycopersicum) as well as in other wild relatives of tomato [START_REF] Kaan | Mise en évidence de la relation entre des résistances de la tomate à deux maladies vasculaires: le flétrissement bactérien (Pseudomonas solanacearum E.F. Sm.) et la Fusariose pathotype 2 (Fusarium oxysporum f.sp. lycopersici (Sacc.) Snyd. & Hans)[END_REF][START_REF] Jaworski | GA 1565-2-4 BWT, GA 219-1-2 BWT, GA 1095-1-4 BWT, and GA 1405-1-2 BWT Bacterial Wilt Tolerant Tomato[END_REF][START_REF] Anaïs | La tomate. Pp[END_REF][START_REF] Mohamed | Genetic nature of bacterial wilt resistance in tomato (Lycopersicon esculentum Mill.) accession LA 1421[END_REF]Carmeille et al. 2006b;[START_REF] Hai | Resistance to Taiwanese race 1 strains of Ralstonia solanacearum in wild tomato germplasm[END_REF]. From these results, it seems that resistance to bacterial wilt is not that frequent in tomato germplasm. The high genetic diversity displayed by Ralstonia solanacearum complex [START_REF] Fegan | Bacterial Wilt: the Disease and the Ralstonia solanacearum species complex[END_REF] and the strong interactions between strains and resistant material [START_REF] Lebeau | Bacterial wilt resistance in tomato, eggplant and pepper: genetic resources challenged with the multifaceted Ralstonia solanacearum species complex[END_REF] suggest that various resistance mechanisms, including strain specific ones, exist in tomato resistant germplasm. Therefore, breeders have some opportunities at their disposal to enlarge the relatively narrow range of resistance sources primarily used so far, and to continue accumulating different mechanisms of resistance in tomato genotypes to obtain better stability of resistance in different environments. However, they might be limited by the fact that some bacterial strains are not controlled by any resistant accession (see section 5. below).

Inheritance studies have focused mostly on F 2 , F 3 and RILs progenies of "Hawaii 7996" crossed with the susceptible "WVa700" (S. pimpinellifolium). Several QTLs of resistance have been identified, including a major QTL on chromosome 6 effective towards "GMI8217", an isolate of race 1 biovar 1 (Thoquet et al. 1996a & b); of "Pss4", an isolate of race 1, biovar 3, phylotype 1 [START_REF] Wang | Worldwide evaluation of an international set of resistance sources to bacterial wilt in tomato[END_REF]; and "JT516", an isolate representative of race 3-phylotype II (Carmeille et al., 2006a). [START_REF] Wang | Resistance of tomato line Hawaii7996 to Ralstonia solanacearum Pss4 in Taiwan is controlled mainly by a major strain-specific locus[END_REF] identified another major QTL of resistance of "Hawaii 7996" effective towards "Pss4" and located on chromosome 12. Several minor QTLs located on chromosomes 3, 4, 8, some of which having a season dependent expression (Carmeille et al., 2006a) have also been identified. Mejia et al (2009) confirmed the QTLs on chromosome 6 and 12 were associated with resistance in "Hawaii 7996" observed in Guatemala field evaluation against local phylotype I strains. Work is ongoing at AVRDC for adding markers to the QTLs regions of "Hawaii 7996" associated to resistance to bacterial strains belonging to phylotype I, in order to develop tools for marker assisted selection.

QTLs of resistance of the resistant line "L285" effective towards "UW364" an isolate of race 1, biovar 4, have also been located on chromosomes 6, as well as on chromosomes 7 and 10 [START_REF] Danesh | Genetic dissection of oligogenic resistance to bacterial wilt in tomato[END_REF]. Pattern of resistance derived from CRA66 has been described as polygenic [START_REF] Prior | Resistance to bacterial wilt (Pseudomonas solanacearum) in tomato: present status and prospects. In: Bacterial Wilt, the disease and its causative agent Pseudomonas solanacearum[END_REF] but no molecular data are available for this source. [START_REF] Kaan | Mise en évidence de la relation entre des résistances de la tomate à deux maladies vasculaires: le flétrissement bactérien (Pseudomonas solanacearum E.F. Sm.) et la Fusariose pathotype 2 (Fusarium oxysporum f.sp. lycopersici (Sacc.) Snyd. & Hans)[END_REF] were the first to mention a quantitative resistance to Fusarium wilt (Fusarium oxysporum f.sp. lycopersici) race 2 in lines bred for bacterial wilt resistance in Puerto Rico, North Carolina, and the French West Indies and they suggested this relation to be more likely of a pleiotropic nature than being due to a genetic linkage. Further, Laterrot & Kaan (1978) as well as Laterrot et al. (1978) have pointed out the frequent association of both these partial resistances with the partial resistance to a third vascular disease, i.e. bacterial canker caused by Clavibacter michiganensis. These authors exemplified this relation between the resistance to the three diseases on a set of varieties bred for bacterial wilt in North Carolina ("NC 72 TR 4-4", "MR4", "Venus" and "Saturn"), in the French West Indies ("Carette", "53 RC", "IRAT L3"), in Burkina Faso ("Farako-Ba"), and in Hawaii ("Hawaii 7996"). They checked that the resistance to Fusarium wilt race 2 observed was not due to the gene I-2. "Kewalo" was the only variety resistant to bacterial wilt that they found susceptible to bacterial canker, and of a low level of resistance to Fusarium wilt race 2. This exception suggests that the mechanisms controlling the resistance to bacterial wilt can be dissociated, in some genotypes, from those involved in the resistance to the two other vascular diseases. Unfortunately Laterrot & Kaan (1978) and Laterrot et al. (1978) did not test bacterial wilt resistant material created in the Philippines and Taiwan, and the general picture of the relationships between the resistance to bacterial wilt and the two other vascular diseases is incomplete.

Resistance to bacterial wilt is sometimes associated to resistance to other bacterial and fungal pathogens

The reciprocal relationship between the resistance to bacterial canker and the two other diseases is verified in some cases, as for instance for "Okitsu Sozai n°1" whose resistance to bacterial canker originates from S. hirsutum var. glabratum PI 134418. This line is also partially resistant to Fusarium wilt race 2 (Laterrot, unpub. results) as well as to some strains of Ralstonia solanacearum [START_REF] Lebeau | Bacterial wilt resistance in tomato, eggplant and pepper: genetic resources challenged with the multifaceted Ralstonia solanacearum species complex[END_REF]. But "Plovdiv 8-12", the resistance of which to bacterial canker originates from a S. pimpinellifolium, is susceptible to Fusarium wilt race 2 (Laterrot et al. 1978); its behaviour towards bacterial wilt is unknown.

The bacterial wilt resistant "Hawaii 7998" was resistant to a race of bacterial spot later confirmed to be race T1 [START_REF] Scott | Sources of resistance to bacterial spot (Xanthomonas campestris pv. vesicatoria (Doidge) Dye) in tomato[END_REF]. Later, Scott et al (1995b) discovered "Hawaii 7981", a line susceptible to race T1, was resistant to bacterial spot race T3 while "Hawaii 7998" was susceptible. Thus, bacterial spot resistance has been found in bacterial wilt resistant lines from Hawaii. For race T1, resistance to bacterial wilt was not correlated with bacterial spot resistance in an F 2 population suggesting separate genes were responsible for resistance to each disease [START_REF] Scott | Bacterial spot resistance is not associated with bacterial wilt resistance in tomato[END_REF]. Resistance to bacterial spot race T4 has been seen in some bacterial wilt resistant breeding lines derived from "Hawaii 7997" even though this line is not resistant to race T4 [START_REF] Scott | Breeding for Bacterial Wilt Resistance in Tomato: The Struggle Continues[END_REF]. The genetic control of this response is not known but again illustrates the association of bacterial wilt resistance with resistance to another disease. A further example is that "Hawaii 7998" had resistance to bacterial canker (Panthee and Gardner, 2010) especially the foliar phase. These examples and those mentioned before indicate the existence of frequent associations between the resistance to some bacteria (vascular or not) and even fungi (vascular) within single tomato genotypes. Hence, [START_REF] Scott | Tomato improvement for bacterial disease resistance for the tropics: A contemporary basis and future prospects[END_REF] advised that when searching for resistance to a given bacterial pathogen, breeders should not overlook genotypes resistant to other bacteria. In this regard, resistance genes can emerge by combining genotypes with some reported bacterial resistance even when one of the parent lines does not show resistance to a particular disease/race. For instance, [START_REF] Hutton | Identification of QTL Associated with Resistance to Bacterial Spot Race T4 in Tomato[END_REF] reported that an allele from "Hawaii 7998" was associated with resistance to bacterial spot race T4 even though this line was susceptible to race T4, suggesting epistasis with genes from the other parent that was T4 resistant. Another possibility is that the pleiotropic nature of resistance in tomato against several diseases could be associated with higher level of expression of systemic acquired resistance (SAR). [START_REF] Lin | Transgenic Tomato Plants Expressing the Arabidopsis NPR1 Gene Confer Enhanced Resistance to A Spectrum of Fungal and Bacterial Diseases[END_REF] found over-expression of Arabidopsis NPR1 (non-expresser of PR genes) gene in a susceptible tomato line could enhance resistance to both Fusarium wilt race 2, bacterial wilt, as well as bacterial spot and gray leaf spot. In this study, they did not test the enhanced resistance against the bacterial canker pathogen.

Lastly, [START_REF] Rouamba | A case of relation between resistances to Pseudomonas solanacearum and Verticillum pathotype 2[END_REF] tested material resistant to bacterial wilt towards a fourth vascular disease, the Verticillium wilt caused by Verticillium race 2, but they found only a loose relationship between both resistances since only two ("IRAT L3" and "Farako-Ba") out of ten lines tested, were resistant to both diseases.

Grafting of susceptible cultivars on resistant rootstocks, an alternative to resistant cultivars

Given the difficulty to create highly resistant lines with good commercial quality, grafting susceptible scions on resistant rootstocks remains an alternative to the cultivation of resistant cultivars. As early as 1969, Gilbert and Chin pointed out that highly resistant tomato lines with poor fruit quality, could be efficiently used as rootstocks on which susceptible scions of good fruit quality could be grafted. These authors reported the bacterial wilt resistance of the root system as being effective even when completely susceptible scions are used. This technique is still used nowadays [START_REF] Cardoso | Potential of Hawaii 7996 hybrid as rootstock for tomato cultivars[END_REF][START_REF] Hanrong | Control of tomato bacterial wilt with grafting[END_REF], though the protection provided by the rootstock is sometimes incomplete [START_REF] Nakaho | Distribution and multiplication of Ralstonia solanacearum in tomato plants with resistance derived from different origins[END_REF]. Indeed, tomato resistant material harbours the bacteria symptomlessly and the resistance is associated with the ability of the plant to restrict bacteria invasiveness [START_REF] Grimault | Some characteristics involved in bacterial wilt (Pseudomonas solanacearum) resistance in tomato[END_REF]. The absence of incompatible interactions in tomato resistant lines (no symptoms, no latent infection) has been confirmed by [START_REF] Lebeau | Bacterial wilt resistance in tomato, eggplant and pepper: genetic resources challenged with the multifaceted Ralstonia solanacearum species complex[END_REF] by testing a core collection of bacterial wilt resistant accessions with a core collection of bacterial strains.

Eggplant is an alternative rootstock for cultivating susceptible tomatoes in contaminated conditions. It was shown to provide a better protection than tomato rootstocks (AVRDC, 1998). This result was confirmed and extended by [START_REF] Lebeau | Bacterial wilt resistance in tomato, eggplant and pepper: genetic resources challenged with the multifaceted Ralstonia solanacearum species complex[END_REF] who found that apart from common cases of latent infection for some eggplant accessions and all tomato accessions, there also exist incompatible interactions between some eggplant resistant lines and some bacterial strains. Further, some eggplant lines control bacterial strains that are not controlled by any of the tomato resistant lines tested so far, as exemplified by Carmeille

Conclusion

The earliest breeding efforts for tomato resistance to bacterial wilt started in Puerto Rico and two American States (North Carolina, Hawaii) 80 years ago followed by programs in the 1950s in Japan and the Philippines. The programs carried out by French research institutes in the Caribbean (and Burkina Faso) started during the course of the 1960s, whereas AVRDC started at the beginning of the 1970s. There were many exchanges of material between the breeding programs for bacterial wilt resistance carried out in several US States, and between them and the Philippines. AVRDC is using and recombining now the resistances bred in the USA, the Philippines and the French West Indies. The breeding material created in Japan was mostly used locally, and perhaps also in Guadeloupe, if one assumes that the Japanese line "OTB2" is equivalent to "CRA66", which is not certain.

Our attempt to draw a general picture of the main sources of resistance to bacterial wilt, of the main breeding programs for this resistance, of their most frequently mentioned resistant varieties, and of the relationships between varieties, is based on a careful work of assembling bits and pieces dispersed in many publications. Given the unavailability of complete information, it is not now possible to come up with a better picture than the one we present here, though more information can emerge out of archives of the scientists, Universities and research institutes involved. Henceforth the synthesis provided here displays the most probable general picture, but it may include some mistakes in the absence of further information.

We added to this survey complementary information on some peculiarities of tomato bacterial wilt resistance. The frequent association of bacterial wilt resistance with resistance to other bacterial or fungal diseases should be of strong interest for breeding and/or research on its genetic basis. Breeding over many decades succeeded in eliminating a number of undesirable traits initially associated to high level of bacterial wilt resistance, but breeding efforts are still ongoing for obtaining large fruit size in resistant material. The adaptation of the breeding material to hot environmental conditions is often mentioned as necessary for obtaining breeding lines which are resistant and agronomically acceptable.

On the whole, we hope this paper to be useful for further research using the plant material mentioned, in particular for comparative genetic studies and breeding concerning the resistance of tomato to bacterial wilt and to other vascular diseases.

Abinne (librarian at INRA-CRAAG, Guadeloupe),and the librarians at North Carolina State University, the University of Hawaii, AVRDC, and the University of Puerto Rico along with Linda Wessel-Beaver for having provided reprints of numerous archives. provide a rationale for pyramiding begomovirus-resistance introgressions into hybrids that will have more durable resistance.

Acronyms found in some names of tomato lines or in

Materials and Methods:

Germplasm: The breeding lines used in these experiments were: Gh13, homozygous for Ty3; Gc143-2, homozygous for Ty1 and Ty3; Gc171, homozygous for Ty3a and Ty4 (provided by J. Scott, University of Florida); CLN2116 homozygous for Ty2 (provided by P. Hanson, The World Vegetable Center) and Gh188-2, also homozygous for Ty2, and Gc21-a, homozygous for Ty3a and lacking Ty4. Gh13 was selected from the hybrid, Favi9 (Vidavsky and Czosnek, 1988). Gc143-2 was selected from a cross of Gc9 (provided by J. Scott, University of Florida) by a susceptible commercial hybrid. Gc21-a was obtained by several cycles of selfing from a cross between Gc171 with a susceptible commercial hybrid and an individual F2 plant. Gh188-2 was selected from a commercial hybrid with Ty2 introgression. The susceptible germplasm, HUJ-VF, without any of the introgressions for resistance, was used in several crosses. HUV-VF was provided by F. Vidavsky, The Hebrew University of Jerusalem. The F2 population used in the segregation of genes Ty3a and Ty4 was obtained from a hybrid produced by a cross between the resistant line Gc171 and a slightly resistant inbred (Gh44). Field Evaluation of disease severity for the different populations: Each entry was replicated three times with 5 plants per replication. Four-week-old seedlings were transplanted into a field near Sanarate, Guatemala, where high levels of viruliferous whiteflies were present. In this area, at least 7 bipartite tomatoinfecting begomoviruses have been identified (Nakhla et al., 2004) as well as the monopartite begomovirus, Tomato yellow leaf curl virus (Mejía and Maxwell, unpublished results). Each plant was scored at about 30-days after transplanting in either January 2009 or October 2009 using a disease severity index (DSI) from zero to six: 0, no symptoms; 1, extremely slight symptoms; 2, slight symptoms; 3, moderate symptoms; 4, severe symptoms with deformed leaves; 5, severe symptoms with stunted plant; and 6, very severe symptoms, no marketable fruit and very stunted plant.

PCR

Development of populations with different introgressions:

Combinations of introgressions Ty3a and Ty4: An F2 population was obtained from a F1 hybrid produced from the cross between line Gc171 (Ty3a/Ty3a, Ty4/Ty4, Scott and Schuster, 2007) by a slightly resistant line (Gh44, ty3/ty3, ty4/ty4). Leaf samples were collected from 77 individual plants for molecular analysis. The genotype of these plants was determined with the PCR primers P6-25F2 and P6-25R5 (SCAR marker P6-25) for introgression Ty3a and a PCR-based marker for Ty4 introgression (Ji et al., 2008). Individual F2 plants were selected that were dominant for one or both genes (RR, Ty3a/Ty3a, Ty4/Ty4; RS, Ty3a/Ty3a, ty4/ty4; SR, ty3/ty3, Ty4/Ty4 and SS, ty3/ty3, ty4/ty4). These plants were selfed to produce F3 seeds. Twelve F3 families, Gc171, Gh44 and a susceptible commercial control C-SS (ty3/ty3, ty4/ty4) were planted in a field to determine their phenotype, i.e., the level of resistance to begomoviruses was determined in January 2009. Each entry was replicated three times with 5 plants per replication.

Combination of introgressions for Ty2 and Ty3: F2 seed was obtained from the cross between resistant line Gh13 (Ty3/Ty3, ty2/ty2, RS-C, Martin et al., 2007), and line CLN2116 (ty3/ty3,Ty2/Ty2, SR-C). The genotype of the F2 plants was determined using molecular markers and individuals with one, both or none of the introgressions for resistance (genotypes RR, Ty3/Ty3,Ty2/Ty2; SR, ty3/ty3, Ty2/Ty2; RS, Ty3/Ty3, ty2/ty2 and SS, ty3/ty3, ty2/ty2) were allowed to self. The genotype of each F3 family was verified and seventeen F3 families were transplanted into a field along with the two parental lines and a susceptible control. Each entry was replicated three times with 5 plants per replication to determine their phenotype and DSIs were taken in October 2009.

Combination of introgressions Ty1, Ty3 and Ty2: F2 seed was obtained from the cross between resistant line Gc143-2 (Ty1/Ty1-Ty3/Ty3, ty2/ty2) and line Gh188-2, (ty1/ty1-ty3/ty3, Ty2/Ty2). The genotype of the F2 plants was determined using molecular markers for Ty3 and Ty2 and individuals with one, both or none of the introgressions were identified (RR, Ty3/Ty3, Ty2/Ty2; RS, Ty3/Ty3, ty2/ty2; SR, ty3/ty3, Ty2/Ty2 and SS, ty3/ty3, ty2/ty2). The Ty1 introgression was not determined as it is linked to Ty3. F3 seed was collected from individual F2 plants of different genotypes and the F3 families were transplanted in the field. Sixteen F3 families and both parental lines were planted and each entry was replicated three times and 5 plants per replication. Plants were scored for their DSIs in October 2009.

Combination of introgressions Ty3a and Ty2: Gc21-a (RS, Ty3a/Ty3a, ty2/ty2), was crossed with line Gh188-2 (SR, ty3/ty3, Ty2/Ty2). Heterozygous individuals were evaluated for resistance (HH, Ty3a/ty3, Ty2/ty2), along with the parental genotypes. Gc21-a (RS) and Gh188-2 (SR) were also crossed to the susceptible line HUJ-VF (ty2/ty2, ty3/ty3). The genotypes of the F1 populations were verified and their resistance phenotype determined in the field in October 2009. Three hybrids [Gc21-a X Gh188-2 (HH, Ty3a/ty3, Ty2/ty2), Gc21-a X HUJ-VF (HS, Ty3a/ty3, ty2/ty2) and Gh188-2 X HUJ-VF (SH, ty3/ty3, Ty2/ty2)], two parental lines and a susceptible commercial control (SS) were planted with three replications of 5 plants each. Plants were scored for their DSIs in October 2009.

Combination of introgressions Ty3a and Ty3: Line Gc21-a (Ty3a/Ty3a) was crossed with line Gh13 (Ty3/Ty3). The level of resistance in the hybrid (Ty3a/Ty3) was evaluated in relation to the parental genotypes. The genotype of the F1 families was determined and subsequently transplanted in the field for the evaluation of their phenotype of resistance. The hybrid (Gc21-a X Gh13), both parental lines, and a susceptible control were planted in with three replications of five plants each. Plants were scored for their DSIs in October 2009.

Results and Discussion:

Natural inoculation with begomoviruses in a field in Guatemala with high populations of the whitefly vector was used to evaluate various combinations of begomovirus-resistance introgressions for their effectiveness to provide resistance to multiple begomoviruses.

Evaluations of the introgressions Ty3a and Ty4:

The highly resistant inbred line, Gc171 (Ty3a/Ty3a, Ty4/Ty4), was crossed with a slightly resistant genotype (Gh44, ty3/ty3, ty4/ty4) (Garcia, et al., 2008a). Among the 77 F2 plants analyzed, 17 were found to be homozygous Ty3a/Ty3a, 12 homozygous ty3a/ty3a, and 48 heterozygous Ty3a/ty3a. With relation to Ty4 introgression, 10 plants were found to be homozygous Ty4/Ty4, 42 homozygous ty4/ty4, and 25 heterozygous Ty4/ty4. Seven F3 families were evaluated: three with genotype ty3/ty3, Ty4/Ty4 (SR), and four with genotype Ty3a/Ty3a, ty4/ty4 (RS) (Fig. 1). Fig. 1. Disease severity index (DSI) and standard errors for F3 families of genotypes RR (4 families Ty3a/Ty3a, Ty4/Ty4), RS (4 families, Ty3a/Ty3a, ty4/ty4), SR (4 families, ty3/ty3, Ty4/Ty4), C-SS (commercial susceptible control), R-RR (parent, Gc171), S-SS (Gh44). The average DSI for the RS F3 families with the Ty3a introgression was 2.1, while the SR F3 families with the Ty4 introgression was 3.6, which is similar to the slightly resistant parent, Gh44 (S-SS). These results indicate that there is a larger effect on resistance by Ty3a than by Ty4 to the multiple bipartite begomoviruses present in this area. This is consistent with the report by Ji et al. (2008) where the Ty4 introgression had a lesser effect on resistance to Tomato yellow leaf curl virus than the Ty3a introgression.

Combinations of introgression Ty2 and Ty3: Seventeen F3 families were obtained from individual F2 plants arising from the cross of Gh13 X CLN2116 (F1, Ty3/ty3, Ty2/ty2). The F3 families were evaluated for symptom development 36 days after transplanting (Fig. 2). Fig. 2. Disease severity index (DSI) with standard errors for the F3 families of genotypes RR (6 families, Ty3/Ty3, Ty2/Ty2), SR (5 families, ty3/ty3, Ty2/Ty2), RS (3 families, Ty3/Ty3, ty2/ty2), SS (3 families, ty3/ty3, ty2/ty2), SR-C (CLN2116, ty3/ty3, Ty2/Ty2), RS-C (Gh13, Ty3/Ty3, ty2/ty2) and SS-C (susceptible commercial hybrid control, ty3/ty3, ty2/ty2). The parents had average DSIs of 0.6 and 3.8 for Gh13 and CLN2116, respectively. A commercial hybrid with neither introgression had a DSI of 5.2. The average DSI of the 6 RR families was 1.6, which was greater than the most resistant parent, Gh13 (DSI 0.6), and was similar to the DSI for the 3 RS families, which had an average DSI of 1.4. The average DSI of the 5 SR families (DSI 2.9) was similar to that for the 3 SS families (DSI 2.6).

These results indicate that Ty3 introgression from Gh13 was mainly responsible for the observed level of virus resistance in the F3 families and adding the Ty2 introgression resulted in no increase in virus resistance. The susceptible control had a DSI of 5.1, while the 3 F3 families without the introgressions (ty3/ty3, ty2/ty2) had a DSI of 2.6, indicating the presence of other unknown genes for resistance.

Of interest was the range of average DSIs within families of one genotype (Fig. 3). For the 6 RR families, the average DSI ranged from 0.8 to 2.3. For the 3 SS families the range was also great, 1.8 to 3.6, for the 3 RS the range was 1.1 to 1.9. For the 5 SR families the range was 2.1 to 3.2. One explanation for these differences among F3 families with the same genotype for the two introgressions could be that there was segregation of other modifying resistance genes in the F2 plants used to generate the F3 families. Combination of introgressions Ty1-Ty3 and Ty2: Individual F2 plants of different genotype obtained from the F1 (Ty1-Ty3/ty1-ty3, Ty2/ty2) of the cross of Gc143-2 X Gh188-2 were selfed to produce 16 F3 families. The DSI for these families was determined in the field (Fig. 4). Fig. 4. Average disease severity index (DSI) and standard errors for 16 F3 families of genotypes RR (5 families, Ty1-Ty3/Ty1-Ty3, Ty2/Ty2), RS (3 families, Ty1-Ty3/Ty1-Ty3, ty2/ty2), SR (3 families, ty1-ty3/ty1-ty3, Ty2/Ty2), SS (5 families, ty1-ty3/ty1-ty3, ty2/ty2), and the parents: RS-C (Gc143-2, Ty1-Ty3/Ty1-Ty3, ty2/ty2) and SR-C (Gh188-2, ty1-ty3/ty1-ty3, Ty2/Ty2). The average DSI for RS families with the T1-Ty3 introgression was 1.7 while DSI for those SR families with the Ty2 introgression was 4.8. RR families with both introgressions for resistance had an average DSI of 1.4 and those with neither introgression had DSI of 3.6. There was no difference in the average DSI for the RR and SR families. Surprisingly, the DSI for the SR families was higher than the DSI for the SS families. These observations indicated that Ty2 introgression provides no effective resistance to these begomoviruses.

An important observation was that there was a considerable range in the average DSI for families with the same genotype for these two introgressions (Fig. 5). For example, one RR family had a DSI of 3.2, which was greater than the other four RR families (range of 0.3 to 1.4). Within the RS families, one family had a DSI of 0.2 and another RS family had a DSI of 3.8. The genotype for the markers in these families, i.e., the presence or absence of the PCR fragments corresponding to the S. lycopersicum size or the introgression size, was reconfirmed by additional testing in the laboratory. Explanations to consider are that there could have been a recombination event between the PCR marker and the resistance gene for these introgressions or that other genes controlling resistance were segregating within the F2 plants. Combinations of genes Ty3a and Ty2: Line Gc21-a (DSI 0.7), homozygous for Ty3a, was crossed to line Gh188-2 (DSI 5.2), homozygous for Ty2; and the DSI for the F1 (HH, Ty3a/ty3, Ty2/ty2) was 0.4. Lines Gc21-a and Gh188-2 were also crossed with susceptible line HUJ-VF (ty3/ty3, ty2/ty2). The resulting heterozygous F1"s, Ty3a/ty3, ty2/ty2 (HS) and ty3/ty3, Ty2/ty2 (SH) had DSIs of 2.9 and 4.2, respectively (Fig. 6). Thus, the Ty2 gene along either in a homozygous or heterozygous condition did not confer an adequate level of resistance. The Ty3a introgression in the heterozygous condition conferred a moderate level of resistance, but when both resistance introgressions were present in the heterozygous condition the F1 was highly resistant. Fig. 6.

Average disease severity index (DSI) and standard errors for the heterozygous genotypes HH (Ty2/ty2, Ty3a/ty3), SH (ty3/ty3, Ty2/ty2), HS (Ty3a/ty3, ty2/ty2) and homozygous parental lines RS (Ty3a/Ty3a, ty2/ty2), SR (ty3/ty3, Ty2/Ty2), and susceptible commercial control SS (ty2/ty2, ty3/ty3). Combinations of introgressions Ty3a and Ty3: Line Gc21-a (Ty3a/Ty3a, DSI 0.3) was crossed to line Gh13 (Ty3/Ty3, DSI 0.4) and DSI for the F1 was 0.2. The average DSI of the commercial susceptible hybrid was 4.0 (Fig. 7). Fig. 7. Average disease severity index (DSI) and standard errors for heterozygous genotypes Gc21-a X Gh13 (Ty3a/Ty3), Gc21-a (Ty3a/Ty3a), Gh13 (Ty3/Ty3) and commercial susceptible hybrid, (com. hybrid).

These results indicate that the combination of introgressions Ty3 and Ty3a confers a high level of resistance (DSI 0.2), and is not different than either of the resistant parents. In a tomato field (about 0.7 ha, at 1,500 m, Department of Sacatepéquez) where symptom incidence was over 90%, samples of young leaves with typical symptoms were collected in December 2009. DNA was extracted at San Carlos University, Guatemala City [START_REF] Garcia | Co-dominant SCAR for detection of the begomovirus-resistance Ty2 locus derived from Solanum habrochaites in tomato germplasm[END_REF]. PCR was performed at North Dakota State tomatoes and peppers grown in the field or greenhouse [START_REF] Brown | First report of "Candidatus Liberibacter psyllaurous" (synonym "Ca. L. solanacearum" associated with "tomato vein-greening" and "tomato psyllid yellows" dieseses in commercial greenhouses in Arizona[END_REF] in Central America.

No universally accepted name exists for this disease on tomatoes, as illustrated by the use of different names: tomato vein-greening in Arizona [START_REF] Brown | First report of "Candidatus Liberibacter psyllaurous" (synonym "Ca. L. solanacearum" associated with "tomato vein-greening" and "tomato psyllid yellows" dieseses in commercial greenhouses in Arizona[END_REF], permanent yellowing in Mexico (Munyaneza et al., 2009b) and Paratrioza disease in Guatemala. At least from our observations in Guatemala several symptoms are notable: flower abortion, purpling of the leaf margins, stem thickening, yellowing of younger leaves, and stunting. An internet search reveals that there is a substantial number of sources using "permanente del tomate" or permanent yellowing disease of tomato. Thus, it is proposed that this diseased be named Liberibacter permanente del tomate or Liberibacter yellowing disease of tomato. This would distinguish this disease from psyllid yellows [START_REF] Brown | First report of "Candidatus Liberibacter psyllaurous" (synonym "Ca. L. solanacearum" associated with "tomato vein-greening" and "tomato psyllid yellows" dieseses in commercial greenhouses in Arizona[END_REF].

Introduction

The phenomenon of cytoplasmic male sterility (CMS) has been described and the genetics underlying the phenomemon studied in many species. Whether arising spontaneously, as the result of mutations, or through alloplasmic incompatibilities in interspecific crosses, the main effect of CMS is on the development of stamens and pollen, leading to aberrant stamens with no pollen or aborted pollen [START_REF] Kaul | Male sterility in higher plants[END_REF]). Other changes correlated with the CMS phenotype are changes in the second whorl affecting petal size and color [START_REF] Andersen | Cytoplasmic sterility in hybrids of Lycopersicon esculentum and Solanum pennellii[END_REF][START_REF] Andersen | Evidence for plasmon differentiation in Lycopersicon[END_REF][START_REF] Petrova | Characterization of cytoplasmic male sterile hybrid line between Lycopersicon peruvianum Mill. x Lycopersicon pennellii Corr. and its crosses with the cultivated tomato[END_REF][START_REF] Farbos | Defective cell proliferation in the floral meristems of alloplasmic plants of Nicotiana tabacum leads to abnormal floral organ development and male sterility[END_REF][START_REF] Leino | Brassica napus lines with rearranged Arabidopsis mitochondria display CMS and a range of developmental aberrations[END_REF] In the tomato, CMS does not occur naturally. CMS has been reported in interspecies hybrids. [START_REF] Andersen | Cytoplasmic sterility in hybrids of Lycopersicon esculentum and Solanum pennellii[END_REF][START_REF] Andersen | Evidence for plasmon differentiation in Lycopersicon[END_REF] reported the emergence and increase of pollen abortion in F1 and backcrosses of the crosses between Solanum lycopersicum, S. cheesmaniiae (formerly L. chesmanii f. typicum and f. minor) or S. habrochaites (formerly L. hirsutum f. glabratum) used as pistillate parents and S. pennellii as the recurrent pollinating parent. Pleiotropic effects of the CMS phenotype included the reduction of anther length and size, and the lengthening of the filaments. The anther size was negatively correlated to the percent of aborted pollen. Similar results were observed by Valkova-Atchkova (1980) in crosses involving S. peruvianum as pistillate parent and S. pennellii and S. habrochaites (formerly L. hirsutum f. typicum) as pollinating parents. Further introgression of the nuclear genome of the recurrent parents confirmed the stability of the CMS phenotype over many generations [START_REF] Petrova | Characterization of cytoplasmic male sterile hybrid line between Lycopersicon peruvianum Mill. x Lycopersicon pennellii Corr. and its crosses with the cultivated tomato[END_REF][START_REF] Stoeva | Towards cytoplasmic male sterility in cultivated tomato[END_REF]).

As a preliminary step to dissecting morphological (and underlying genetic) changes occurring in the CMS phenotype, this study has focused on the comparative analysis of the size of epidermal cells from abaxial and adaxial sides of petals and stamens of mature flowers from CMS-pennellii line (0% stainable pollen), the isonuclear S. pennellii (100% pollen fertility), and the cultivated tomato S. lycopersicum.

Materials and methods

Plant material

In the study the following genotypes were used: Solanum lycopersicum (cv. Merkurii), Solanum pennellii (LA716), and CMS-pennellii (CMS line) previously described in Petrova et al. (1998) and [START_REF] Radkova | Morphological, cytogenetic and molecular genetic studies of cytoplasmic male sterility in genus Lycopersicon[END_REF]. To determine the size of the epidermal cells, fully expanded flowers in stage 20 according to the classification of [START_REF] Burkhin | Flower development schedule in tomato Lycopersicon esculentum cv. sweet cherry[END_REF] were collected from plants grown in the same environmental chamber with 24 C/ 20 C day/night temperature and 18/6 hours day/night photoperiod (Fig. 1).

Tomato flower preparation for confocal microscopy:

A 0.1% aqueous solution of calcofluor white (SIGMA) was made [START_REF] Pringle | Staining of bud scars and other cell wall chitin with calcofluor[END_REF]). Excess solution was kept in the freezer and defrosted as needed. Excised floral structures (petals, anthers or filaments) were immersed in approximately 5 ml of 0.1% calcofluor solution and placed under vacuum until rapid boil was achieved. The sunken specimens were allowed to soak for 20 minutes in the dark and then were rinsed in dH 2 O for 10 minutes. Specimens were dissected and placed on slides. For the anthers, spacers of approximately 0.5 mm -1 mm thick were used to ensure the specimens were not smashed by the coverslip. Coverslips were placed on top and fixed with nail polish at each corner. Calcofluor staining of epidermal cell walls was observed using the DAPI channel of the confocal laser-scanning microscope (CLSM) (FV1000). Images were taken at the planes in which cell wall margins contacted each other. Cells were counted in image frames measuring from 150 m X 150 m to 300 m X 270 m, with the frame size adjusted as needed to include from five to twenty cells in each frame.

Ten to fifteen cells from both sides (abaxial and adaxial) of each floral structure were measured using ImageJ [START_REF] Rasband | [END_REF] to obtain cell areas (in m 2 ). The ANOVA and Tukey's LSD statistical tests (Minitab, Inc.) were performed separately for each epidermal floral surface across genotypes to determine if any significant difference existed between the mean surface areas of studied epidermal cells.

Results

Petal abaxial cell surface area was significantly different among the three genotypes (p < 0.001) with CMS line having the largest mean cell size and S. pennellii having the smallest (Figs. 2,3). Petal adaxial cell sizes differed significantly with S. lycopersicum and S. pennelli both being larger than the CMS line (Fig 3). Although no statistical test was done, abaxial cells were smaller than the adaxial in the two species S. pennellii and S. lycopersicum. On the contrary, the abaxial cells in the CMS line were larger in comparison to the adaxial cells.

Epidermal cell area on both the adaxial and the abaxial surfaces of the anthers was significantly different (p< 0.001) among the three genotypes (Figs. 4,5), with S. lycopersicum having the largest mean cell size for both, and the CMS line, the smallest.

Among the filament epidermal cells, the abaxial cells in the CMS line and cultivated tomato were not significantly different, but both were significantly bigger than those in S. pennellii (p< 0.001) (Fig. 6,7). There were no significant differences in the mean size of the epidermal cells on the adaxial surface of the filaments of the three strains (p=0.252) (Fig. 5,6). There was a large variation (Fig. 7) in the size of the adaxial epidermal cells in S. pennellii and the CMS line, while the size of the adaxial cells of filament in the cultivated tomato showed less variation.

Discussion

According to published data, cytoplasmic male sterility has a considerable effect on the size and color of petals and stamens [START_REF] Kaul | Male sterility in higher plants[END_REF][START_REF] Farbos | Defective cell proliferation in the floral meristems of alloplasmic plants of Nicotiana tabacum leads to abnormal floral organ development and male sterility[END_REF][START_REF] Leino | Brassica napus lines with rearranged Arabidopsis mitochondria display CMS and a range of developmental aberrations[END_REF]. Our previous studies had also shown that in comparison to S. pennellii, the CMS flowers have smaller and lighter green petals and anthers and longer filaments (Fig 1; Petrova et al 1998). In addition to tomato nuclear male sterility mutants (reviewed in Gorman and McCormick 1997) ), similar changes of flower structures have been reported for tomato plants grown at low temperatures [START_REF] Lozano | Tomato flower abnormalities induced by low temperatures are associated with changes of expression of MADS-box genes[END_REF]) and in lossof-function transgenic plants for the B-class genes required for petal and stamen identity [START_REF] De Martino | Functional analysis of two tomato APETALA3 genes demonstrate diversification in their roles in regulating floral development[END_REF].

The size of plant organs is determined by cell division and cell expansion and elongation. Our investigation of the size of epidermal cells shows that the smaller size of the anthers in the CMS line (Fig. 1; Stoeva-Popova et al., unpublished) can be explained largely by the reduction in the size of the cells. The epidermal cells on both surfaces of the CMS anther were significantly smaller in comparison to S. pennellii: respectively 51% smaller on the adaxial side, and 42.3% on the abaxial surface. On the other hand, no corresponding effect of cytoplasmic male sterility on cell size in the petals was observed On the abaxial surface of the petals of CMS line, the cells were 81.5% larger than in S. pennellii, while on the adaxial side, they were 32.2 % smaller. According to our measurements (Stoeva et al., unpublished data) the CMS-pennellii filaments are several magnitudes longer than the filaments of S. pennellii (Fig. 1). This is not merely a consequence of increased cells size, as the filament epidermal cells of CMS are statistically larger than those of S. pennellii only on the abaxial side. CMSpennellii and S. pennellii share the same nuclear genome and cytoplasmic male sterility affects expression of nuclear genes ( reviewed in Linke, Börner 2005, Chase 2006). Our results above show that cytoplasmic male sterility does not equally affect the development of petals, stamens and filaments.

Our study has shown that there are significant differences in the size of epidermal cells between the two species: the red-fruited cultivated tomato and the green-fruited S. pennellii. Significant differences were determined for the epidermal cells of the anthers and petals. The epidermal cells of the anthers of the cultivated tomato were larger, which is an indication that the anthers of S. pennellii have greater number of epidermal cells. The same conclusion could be drawn from the study of the abaxial epidermal cells of the petals and the filaments. Since the two species are not closely related it will be interesting to investigate other representatives of the tomato clade and to determine if the size of epidermal cells of flower structures can be indicative of species relatedness.

Although no statistical analysis was carried out, our data show one consistent feature across all genotypes: the largest epidermal cells were observed on anthers, while the ones with the smallest surface area were epidermal cells of petals. The following list of 1,196 accessions of wild tomatoes and allied Solanum species is a revision of the list published in TGC vol. [START_REF] Lebeau | Bacterial wilt resistance in tomato, eggplant and pepper: genetic resources challenged with the multifaceted Ralstonia solanacearum species complex[END_REF]2007. Other types of TGRC stocks are catalogued in TGC 58 (monogenic mutants) and TGC 59 (miscellaneous stocks). Inactive accessions have been dropped and new collections added to the present list. The new material includes populations of wild or feral cherry tomato (S. lycopersicum 'cerasiforme") from Mexico (LA4352, LA4353), a stock of S. pimpinellifolium containing the sun gene introgressed from S. lycopersicum, and populations of S. peruvianum from the Azapa valley of northern Chile (LA4445-LA4448).

Seed samples will be provided, upon request, for research, breeding or educational purposes. Some accessions may be temporarily unavailable for distribution during seed multiplication. In general, only small quantities of seed will be provided: 25 seed per accession for the self-pollinated accessions, 50 for the outcrossers or facultative accessions, and 5-10 for the allied Solanum species. These seed samples should be sufficient for researchers to produce larger quantities of seed, if needed. Accessions are grown for seed increase in the UC-Davis greenhouses, except for cherry tomatoes and certain populations of S. pimpinellifolium, which are grown in the field. The population sizes used for seed multiplication depend on the mating system, and are designed to maintain genetic diversity within accessions (see guidelines at http://tgrc.ucdavis.edu).

The following tables are ordered by species name, using the classification system of Peralta et al. ( 2008) 11 , but with the equivalent Lycopersicon names listed as well. Although only brief collection site data can be presented here, more detailed records are available from our website, including geographic coordinates, images, and donor information. An appendix table lists the accessions belonging to the core subsets for each species.

S. arcanum

Fig 2 - 12 :

 212 Fig 2-12: Pedigree Montage Research Reports Preliminary Observations on the Effectiveness of Five Introgressions for Resistance to Begomoviruses in Tomatoes Luis Mejía, Rudy E. Teni, Brenda E. García, Ana Cristina Fulladolsa, and Luis Méndez; Sergio Melgar, and Douglas P. Maxwell Preliminary report on association of 'Candidatus Liberibacter solanacearum' with field grown tomatoes in Guatemala Luis Mejía, Amilcar Sánchez, and Luis Méndez; D. P. Maxwell; R. L. Gilberston; V.V. Rivera and G.A. Secor Study of epidermal cell size of petals and stamens in tomato species and hybrids using confocal laser-scanning microscopy Christopher Lofty, Julian Smith, Pravda Stoeva-Popova

  et al. (2006b), Wicker et al. (2007) and Lebeau et al. (2011).

Figure 1 :

 1 Figure 1: General scheme of the relationships between worldwide programs and lines. Notations in color correspond to Literature Cited starting on page 23.

Figure 3 :Figure 9 :

 39 Figure 3: Pedigree of 'MR4' & 'NC 72 TR 4-4' (and of 'Venus' and 'Saturn') (North Carolina State University material) (Taken from Daunay, 1977, based on W.R. Henderson & E. Echandi, pers. com. to H. Laterrot in the 1970s, and on Hendersons & Jenkins, 1972)

  Methods for detection of the introgressions associated with resistance to begomoviruses: DNA extraction was as reported by García et al. (2008). PCRbased molecular markers have been developed for the detection of Ty2, Ty3, Ty3a, and Ty4. Garcia et al. (2007) developed a co-dominant SCAR (Sequence Characterized Amplified Region) marker from the RFLP T0302 marker at 89 cM for the detection of the Ty2 introgression. The Ty3 and Ty3a introgressions were monitored with the co-dominant SCAR marker P6-25 (Ji et al., 2007a). The Ty4 introgression was detected with PCR primers developed by Y. Ji and J. Scott (personal communication, Ji et al., 2008). The presence of the Ty1 introgression in Gc143-2 was determined by sequencing the PCR fragments associated with the RFLP TG97 marker (García and Maxwell, unpublished).

Fig. 3 .

 3 Fig. 3. Disease severity index (DSI) and standard errors for the individual F3 families with the genotypes as listed in Fig. 2 along with the parents and susceptible commercial control.

Fig. 5 .

 5 Fig. 5. Average disease severity index (DSI) and standard errors for individual F3 families for each genotype and the two parents. See Fig. 3 for codes.

  of tomatoes has received considerable attention in the local newspapers in Guatemala. Locally, it is referred to "Paratrioza disease", which refers to the insect associated with symptomatic plants. The symptoms on tomatoes are flower abortion, purple margins of youngest leaves, upward cupping of leaves, thickened stems and retarded internode growth, and stunting of the plants (Fig.1). In Mexico, a disease of tomato with similar symptoms is called permanent damage disease or permanent yellowing disease (daňo permanente del tomate; Páramo Menchaca, 2007) andMunyaneza et al. (2009) reported that Candidatus Liberibacter solanacearum was associated with these plants. This unculturable bacterium is transmitted by the tomato/potato psyllid (Bactericerca (Paratrioza) cockerelli).

Fig. 1 .

 1 Fig. 1. Typical symptoms associated with "Paratrioza disease" of tomato in Guatemala. Image taken December 2009 and shows flower abortion, purple leaf margins, cupping of leaves and thickened petioles and stems.

Fig 2 :Fig 3 :Fig. 4 :Fig. 5 :Fig. 6 :Fig 7 :

 234567 Fig 2: CLSM photographs of epidermal cells from abaxial and adaxial surfaces of petals from flowers (genotypes and magnification as indicated on pictures)

Table of Contents Foreword Announcements Feature Article Tomato resistance to bacterial wilt caused by Ralstonia solanaearum E.F. Smith: ancestry and peculiarities

  Daunay M.C., Laterrot H., Scott J.W., Hanson P., Wang J.-F

	Fig 1: Origins Tomato Bacterial Wilt material

Table 1 :

 1 Summing up of the phenotype of some breeding lines

Table 1 . Summing up of the phenotype of some breeding lines created or used in the breeding programmes of North Carolina, Hawaii, Florida, Japan, Philippines, French West Indies, and Taiwan.

 1

		line or
		accession				fruit
	origin	name	growth habit	fruit shape	fruit size	colour	source
							Laterrot et al.
							(1978); INRA
	North Carolina			slightly			germplasm
	University	NC 72 TR 4-4 indeterminate	flattened	60-100 g	red	database
	North Carolina						Laterrot et al.
	University	MR4	indeterminate				(1978)
	North Carolina						Suzuki et al.
	University	NC1953-60N			6,5 g		(1964)
	North Carolina						Suzuki et al.
	University	NC19/53-64N			7,6 g		(1964)
							Kaan et al.
							(1975); INRA
							germplasm
							database;
	North Carolina						Henderson &
	University	Saturn	indeterminate	deep globe	100-140 g	red	Jenkins (1972)
							Laterrot et al.
							(1978); INRA
							germplasm
							database;
	North Carolina						Henderson &
	University	Venus	indeterminate	slightly oblate	> 180 g	red	Jenkins (1972)
	Hawaii
	University	HES 5808-2	indeterminate		15g
							INRA germplasm
	Hawaii						database, Scott.
	University	H7996	determinate	small oblate	20-60g	red	pers. com.
							Wang et al.
							(1998), Scott et
	Hawaii						al. (2005), Scott.
	University	H7997	indeterminate	small oblate	30-80 g	red	pers. com.
							Wang et al.
							(1998), Scott et
	Hawaii						al. (2005), Scott.
	University	H7998	indeterminate	small oblate	30 g	red	pers. com.
	Hawaii						INRA germplasm
	University	Kewalo	determinate	flattened	140-180 g	red	database
	University of						Scott et al.
	Florida	Neptune	determinate		123-136g	red	(1995a)
	Hort. Res.						Wang et al.
	Sta.,Japan	BF-Okitsu	indeterminate		15-20 g	red	(1998)
	Hort. Res.						Suzuki et al.
	Sta.,Japan	OTB1			13,8 g	pink	(1964)
	Hort. Res.						Suzuki et al.
	Sta.,Japan	OTB2			30,6 g	red	(1964)

  (L. peruvianum or L. peruvianum var. humifusum)

	S. arcanum (L. peruvianum or L. peruvianum var. humifusum) S. galapagense S. lyc. cerasiforme S. lyc. cerasiforme S. pimpinellifolium SolCAP SolCAP
	LA1031 LA0456 LA2767 LA4117B San Pedro -Paso Jama Balsas Clemesi Chitita LA1681 Mushka LA2128 Zumbi LA1537 Azapa Valley LA1259 Catarama LA1562 Cieneguilla LA1615 Piura to Simbala LA1925 Pangaravi LA2578 Tuturo LA3798 Río Pativilca LA1137 LA4133 LA0422		Amazonas Moquegua Tarapaca Antofagasta Lima Zamora-Chinchipe Tarapaca Los Rios Lima Piura Ica Ancash Ancash LA1206 LA0411 LA1654	Peru Peru Chile Chile Peru Ecuador Chile Ecuador Peru Peru Peru Peru Peru
	LA1032 LA0458 LA2768 LA4118 LA1691 LA2144 LA1556 LA1260 LA1571 LA1617 LA1933 LA2585 LA3803 LA1141 LA0247 LA0438	Aricapampa Tacna Empalme Codpa Toconao Yauyos Chanchan Hacienda Higuereta Pueblo Viejo San Jose de Palle Tumbes South Jaqui (4x, origin unknown) Pacanguilla		La Libertad Tacna Tarapaca Antofagasta Lima Chimborazo Lima Los Rios Lima Tumbes Arequipa La Libertad LA1228 LA0417 LA1656	Peru Peru Chile Chile Peru Ecuador Peru Ecuador Peru Peru Peru Peru
	LA1346 LA0460 LA2771 LA4119 LA1695 LA2155 LA1616 LA1261 LA1572 LA1618 LA1936 LA2628 LA3852 LA1401 S. neorickii LA0446	Casmiche Palca Above Poconchile Socaire Cacachuhuasiin, Cacra Maydasbamba La Rinconada Babahoyo Hacienda Huampani Tumbes North Huancalpa Echarate Atinchik, Pachacamac		La Libertad Tacna Tarapaca Antofagasta Lima Cajamarca Lima Los Rios Lima Tumbes Arequipa Cusco Lima LA1231 LA0442 LA1668	Peru Peru Chile Chile Peru Peru Peru Ecuador Peru Peru Peru Peru Peru
	LA1350 LA0470 LA2773 LA4120 LA1696 LA2156 LA1675 LA1262 LA1573 LA1628 LA1950 LA2645 LA3859 LA1410 LA1319 LA0716	Chauna Taltal Zapahuira Cahuisa Huanchuy to Cacra Ingenio Montan Toparilla Canyon Milagro Empalme Nana Huanchaco Pescadores Desvio Chulucanas-Morropon TYLCV resistant selection "hirsute"		Cajamarca Antofagasta Tarapaca Tarapaca Lima Cajamarca Lima Guayas Lima La Libertad Arequipa Piura LA1268 LA1237 LA1674	Peru Chile Chile Chile Peru Peru Peru Ecuador Peru Peru Peru Peru
	LA1351 LA1029 LA2774 LA4121 LA1717 LA2158 LA1692 LA1263 LA1575 LA1629 LA1987 LA2646 LA3910 S. habrochaites Rupe Moquegua Socorama Pachica -Poroma Sopalache Rio Chotano Putinza Barranco Chico Huaycan Barrancos de Miraflores Viru-Fundo Luis Enrique Chalaco Near tortoise preserve, Santa Cruz LA1322 LA0722		Cajamarca Moquegua Tarapaca Tarapaca Piura Cajamarca Lima Guayas Lima Lima La Libertad Piura Galapagos Islands LA1286 LA1245 LA1701	Peru Peru Chile Chile Peru Peru Peru Ecuador Peru Peru Peru Peru Ecuador
	LA1360 LA1030 LA2778 LA4122 LA1718 LA2159 LA1913 LA1269 LA1576 LA1630 LA1992 LA2647 LA4027 LA0407 LA1626A LA1028	Pariacoto Tarata Rd. Chapiquina Chiapa Huancabamba Atonpampa Tinguiayog Pisiquillo Manchay Alta Fundo La Palma Pichicato Morropon-Chalaco Olmos-Jaen Road		Ancash Tacna Tarapaca Tarapaca Piura Cajamarca Ica Lima Lima Ica Lima Piura Lambayeque LA1307 LA1246 LA1712	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Peru Peru Peru
	LA1394 LA1782 LA2779 LA4127 LA1721 LA2167 LA1929 LA1279 LA1577 LA1631 LA1993 LA2652 LA4138 LA1223 LA1716 LA1037	Balsas -Rio Utcubamba Quebrada de Acari Cimentario Belen Alto Umayani Ticrapo Viejo Cimentario Cajamarca La Yapana (Rio Ingenio) Cieneguilla Cartavio Planta Envasadora San Fernando (Moche) Chicama Valley? Sullana El Corregidor, La Molina	Amazonas Arequipa Tarapaca Tarapaca Huancavelica Cajamarca Ica Lima La Libertad La Libertad Lima Piura Lima LA1314 LA1261 LA1732	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Peru Peru Peru
	LA1395 LA1917 LA2780 LA4129 LA1731 LA2171 LA1935 LA1280 LA1578 LA1633 LA2093 LA2653 LA1266 LA2113 LA1141	Chachapoyas Llauta (4x) Belen to Lupica Pachica (Rio Camarones) Rio San Juan El Molino Lomas de Atiquipa Chontay Santa Marta Co-op Huayna Capac La Union San Francisco de Chocon Querecotillo		Amazonas Ayacucho Tarapaca Tarapaca Huancavelica Piura Arequipa Lima La Libertad Ica El Oro Piura LA1320 LA1279 LA1777	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Ecuador Peru
	LA1396 LA1930 LA2879 LA4132 LA1736 LA2174 LA1947 LA1301 LA1579 LA1634 LA2096 LA2655 LA1347 LA2133 LA1208	Balsas (Chachapoyas) Quebrada Calapampa Peine Esquina Pucutay Rio Chinchipe, San Augustin Puerto Atico Hacienda San Ignacio Colegio Punto Cuatro #1 Fundo Bogotalla #1 Playa La Huaca to Sullana S. sitiens (S. rickii) Amazonas Arequipa Antofagasta Tarapaca Piura Cajamarca Arequipa Ica Lambayeque Ica Loja Piura LA1323 LA1301 LA1809	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Ecuador Peru
	LA1626 LA1932 LA2880 LA4319 LA1737 LA2175 LA1949 LA1332 LA1580 LA1635 LA2097 LA2656 LA1974 LA1353 LA2190 LA1237	Mouth of Rio Rupac Minas de Acari Quebrada Tilopozo Alto Rio Lluta Cashacoto Timbaruca Las Calaveritas Nazca Colegio Punto Cuatro #2 Fundo Bogotalla #2 Macara Suarez Chuquicamata	Ancash Arequipa Antofagasta Tarapaca Piura Cajamarca Arequipa Ica Lambayeque Ica Loja Tumbes Antofagasta LA1338 LA1335 LA1912	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Ecuador Peru Chile
	LA1708 LA1938 LA2882 LA4321 LA1738 LA2196 LA1951 LA1335 LA1581 LA1636 LA2102 LA2659 LA2876 LA1361 LA2198 LA1246	Chamaya to Jaen Quebrada Salsipuedes Camar Quebrada Cardones Desfiladero Caclic Ocona Pescaderos Punto Cuatro Laran El Lucero Castilla, Univ. Nac. de Piura Chuquicamata	Cajamarca Arequipa Antofagasta Tarapaca Piura Amazonas Arequipa Arequipa Lambayeque Ica Loja Piura Antofagasta LA1385 LA1371 LA1926	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Ecuador Peru Chile
	LA1984 LA1958 LA2884 LA4324 LA1739 LA2204 LA1954 LA1341 LA1582 LA1637 LA2112 LA2718 LA2877 LA1363 LA2319 LA1269	Otuzco Pampa de la Clemesi Ayaviri Estacion Puquio Canchaque to Cerran Balsapata Mollendo Huampani Motupe La Calera Hacienda Monterrey Chilca El Crucero	La Libertad Moquegua Antofagasta Tarapaca Piura Amazonas Arequipa Lima Lambayeque Ica Loja Lima Antofagasta LA1388 LA1375 LA1930	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Ecuador Peru Chile
	LA1985 LA1959 LA2887 LA4327 LA1740 LA2314 LA1955 LA1342 LA1583 LA1638 LA2145 LA2725 LA2878 LA1559 LA2325 LA1272	Casmiche Huaico Moquegua Quebrada Bandurria Pachica, Rio Camarones Huancabamba San Francisco Matarani Casma Tierra de la Vieja Fundo El Portillo Juan Montalvo Tambo Colorado Mina La Exotica	La Libertad Moquegua Antofagasta Tarapaca Piura Amazonas Arequipa Ancash Lambayeque Lima Los Rios Ica Antofagasta LA1420 LA1478 LA1941	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Ecuador Peru Chile
	LA2150 LA1960 LA2888 LA4329 LA1741 LA2321 LA1975 LA1343 LA1584 LA1645 LA2146 LA2805 LA2885 LA1624 S. pennellii LA1274	Puente Muyuno (Rio Jequetepeque) Rio Osmore Loma Paposo Puente del Diablo, Rio Salado Sondorilla Chirico Desvio Santo Domingo Puente Chao Jayanca to La Vina Banos de Miraflores Hacienda Limoncarro cv. Indehiscent Currant Caracoles	Cajamarca Moquegua Antofagasta Antofagasta Piura Amazonas Lima La Libertad Lambayeque Lima La Libertad Antofagasta LA1425 LA1521 LA1946	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Peru Chile
	LA2151 LA1961 LA2891 LA4330 LA1753 LA2324 LA1977 LA1344 LA1585 LA1651 LA2147 LA2831 LA4105 LA1718 LA0716 LA1282	Morochupa (Rio Jequetepeque) Toquepala Taltal Caspana Surco Leimebamba Orcocoto Laredo Cuculi Vivero, La Molina Yube Rio Nazca Mina La Escondida	Cajamarca Tacna Antofagasta Antofagasta Lima Amazonas Lima La Libertad Lambayeque Lima Cajamarca Ica Antofagasta LA1429 LA1547 LA1953	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Peru Peru Chile
	LA2152 LA1963 LA2930 LA4332 LA1764 LA2329 LA1989 LA1345 LA1586 LA1652 LA2149 LA2832 LA4110 LA1721 LA0751 LA1283	San Juan #1 (Rio Jequetepeque) Rio Caplina Quebrada Taltal Rio Grande West of Canta Aricapampa (self-fertile selection, origin unknown) Samne Zana, San Nicolas Cieneguilla Puente Muyuno Chicchi Tara Mina San Juan	Cajamarca Tacna Antofagasta Antofagasta Lima La Libertad La Libertad La Libertad Lima Cajamarca Ica Antofagasta LA1453 LA1576 LA1963	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Peru Chile
	LA2153 LA1965 LA2931 LA4334 LA1772 LA2409 LA2573 LA1348 LA1587 LA1659 LA2170 LA2833 LA4112 LA1731 LA1272 LA1290	San Juan #2 (Rio Jequetepeque) Causuri Guatacondo Quebrada Sicipo West of Canta Miraflores Valle de Casma Pacasmayo San Pedro de Lloc Pariacoto Pai Pai Hacienda Asiento Aguada Limon Verde	Cajamarca Tacna Tarapaca Antofagasta Lima Lima Ancash La Libertad La Libertad Ancash Cajamarca Ica Antofagasta LA1456 LA1578 LA1973	Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Peru Peru Chile
	LA2157 LA1967 LA2932 LA4335 LA1775 LA2552 LA2581 LA1349 LA1588 LA1660 LA2173 LA2836 LA4113 LA1753 LA1277 LA1301	Tunel Chotano Pachia, Rio Caplina Quebrada Gatico, Mina Escalera Quebrada Tucuraro Rio Casma Las Flores Chacarilla (4x) Cuculi Laredo to Barraza Yautan to Pariacoto Cruz de Huayquillo Fundo Pongo Estacion Cere	Cajamarca Tacna Antofagasta Antofagasta Ancash Cajamarca Tarapaca Lambayeque La Libertad Ancash Cajamarca Ica Antofagasta LA1461 LA1582 LA2076	Peru Peru Chile Chile Peru Peru Chile Peru Peru Peru Peru Peru Chile
	LA2163 LA2164 LA2172 LA2185 LA1968 LA1969 LA1970 LA1971 LA2946 LA2949 LA2952 LA2955 LA4336 LA1777 LA2556 LA2732 LA1355 LA1589 LA1661 LA2176 LA2839 LA4114 LA1777 LA1356 LA1314 LA4337 LA1778 LA2567 LA2742 LA1357 LA1590 LA1670 LA2178 LA2840 LA4115 LA1918 LA1367 LA1322 LA4338 LA4339 LA1779 LA1918 LA2574 LA2648 LA2744 LA2745 LA1359 LA1370 LA1591 LA1592 LA1676 LA1678 LA2179 LA2180 LA2850 LA4116 LA1928 LA1376 LA1331 LA2851 LA4331 LA2098 LA1656 LA1338	Cochabamba to Yamaluc Yamaluc Cuyca Pongo de Rentema Cause Seco Estique Pampa Tarata Palquilla Guatacondo Chusmisa Camiña Quistagama Quebrada Cascabeles Rio Casma Puente Moche Moquella Nepena Viru to Galunga Esquina de Asia Timbaruca Tialango Pampa Carbonatera Quebrada Paposo Rio Casma Quita Camarones-Guancarane Jimbe Viru to Tomaval Rio Sama Tororume San Hilarion de Tomaque Quebrada desde Cerro Oeste de Paqui Antofagasta Cajamarca Cajamarca Cajamarca Amazonas Tacna Tacna Tacna Tacna Tarapaca Tarapaca Tarapaca Tarapaca Antofagasta Ancash La Libertad Tarapaca Ancash La Libertad Lima Cajamarca Amazonas Antofagasta LA1464 LA1584 LA2077 Antofagasta Ancash Ancash Tarapaca Ancash La Libertad Tacna Cajamarca Amazonas LA1482 LA1586 LA2078 Quebrada Taltal, Estacion Breas Antofagasta Quebrada Los Zanjones Antofagasta Rio Casma Ancash Llauta Ayacucho Cullaspungro Ancash Santo Domingo Piura Sobraya Tarapaca Pan de Azucar Tarapaca La Crau Ancash San Jose de Palla Lima Ascope La Libertad Moche La Libertad Fundo Huadquina, Topara Ica San Juan Lucumo de Topara Ica Tamboripa -La Manga Cajamarca La Coipa Cajamarca Santa Rosa, Manta Manabi Quebrada de Paqui Antofagasta LA1483 LA1590 LA2093 La Carcel de Montecristo Manabi Cerro Quimal Antofagasta LA1509 LA1593 LA2099	Peru Peru Peru Peru Peru Peru Peru Peru Chile Chile Chile Chile Peru Peru Chile Peru Peru Peru Peru Peru Chile Chile Peru Peru Chile Peru Peru Peru Peru Peru Chile Chile Chile Chile Peru Peru Peru Peru Chile Chile Peru Peru Peru Peru Peru Peru Peru Peru Chile Ecuador Ecuador Chile
	LA2326 LA1972 LA2980 LA1927 LA2650 LA2770 LA1371 LA1593 LA1679 LA2181 LA2852 LA2103 LA1674 LA1340	Above Balsas Rio Sama Yacango Ocobamba Ayabaca Lluta Santa Eulalia Puente Chao Tambo de Mora Balsa Huaico Cirsto Rey de Charapoto		Amazonas Tacna Moquegua Ayacucho Piura Tarapaca Lima La Libertad Ica Cajamarca Manabi LA1511 LA1599 LA2126A	Peru Peru Peru Peru Peru Chile Peru Peru Peru Peru Ecuador
	LA2327 LA2404 LA2981A Torata to Chilligua en route to Puno Aguas Calientes Arica to Tignamar S. chmielewskii (L. chmielewskii) Cajamarca Tarapaca Moquegua LA1928 Ocana Ayacucho LA2651 Puente Tordopa Piura LA2834 Hacienda Asiento Ica LA1374 Ingenio Ica LA1594 Cerro Sechin Ancash LA1680 La Encanada Lima LA2182 Cumba Amazonas LA2853 Experiment Station, Portoviejo-INIAP Manabi LA2109 LA1542 LA1724 LA1602 LA1346 LA2131	Peru Chile Peru Peru Peru Peru Peru Peru Peru Peru Ecuador
	LA2328 LA2405 LA3111 LA1028 LA1978 LA2722 LA2955B Quistagama Aricapampa Tignamar Tarata Casinchihua Colca Puente Auco LA1375 San Vicente de Canete LA1595 Nepena to Samanco LA1682 Montalban -San Vicente LA2183 Corral Quemado LA2854 Jipijapa LA2119 LA1732 LA1406		La Libertad Tarapaca Tacna Apurimac Ancash Lima Tarapaca Lima Ancash Lima Amazonas Manabi LA1543 LA1606 LA2135	Peru Chile Peru Peru Peru Peru Chile Peru Peru Peru Peru Ecuador
	LA2330 LA2406 LA3112 LA1306 LA2092 LA2812 LA2959 LA1380 LA1596 LA1683 LA2184 LA2857 LA2128 LA1733 LA1455	Chagual Arica to Putre Estique Pampa Tambo Chinuko Lambayeque Chaca to Caleta Vitor Chanchape Santa to La Rinconada Miramar Bagua Isabela: Puerto Villamil		La Libertad Tarapaca Tacna Ayacucho Chimborazo Lambayeque Tarapaca Piura Ancash Piura Amazonas Galapagos Islands LA1620 LA1617 LA2137	Peru Chile Peru Peru Ecuador Peru Chile Peru Peru Peru Peru Ecuador
	LA2331 LA2731 LA3113 LA1316 LA2098 LA2855 LA2964 LA1381 LA1597 LA1684 LA2186 LA2866 LA2155 LA1926 LA1457	Agallapampa Moquella Apacheta Ocros Sabianga Mollinomuna, Celica Quebrada de Burros Naupe Rio Casma Chulucanas El Salao Via a Amaluza		La Libertad Tarapaca Tacna Ayacucho Loja Loja Tacna Lambayeque Ancash Piura Amazonas Loja LA1622 LA1659 LA2163	Peru Chile Peru Peru Ecuador Ecuador Peru Peru Peru Peru Peru Ecuador
	LA2333 LA2737 LA3114 LA1317 LA2099 LA2860 LA3218 LA1382 LA1598 LA1685 LA2187 LA2914A Urb. La Castellana, Surco Casmiche Yala-yala Quilla Hacienda Pajonal Sabiango to Zozoranga Cariamanga Quebrada Guerrero (Islay) Chachapoyas to Balsas Culebras to La Victoria Marcavelica La Caldera LA2158 LA1946 LA1464		La Libertad Tarapaca Tacna Ayacucho Loja Loja Arequipa Amazonas Ancash Piura Amazonas Lima LA2078 LA1683 LA2181	Peru Chile Peru Peru Ecuador Ecuador Peru Peru Peru Peru Peru Peru
	LA2334 LA2739 LA3115 LA1318 LA2100 LA2861 LA3220 LA1383 LA1599 LA1686 LA2188 LA2914B La Castellana, Surco San Juan Nama to Camina W of Quilla Auquibamba Sozorango Las Juntas Cocachacra Chachapoyas to Bagua Huarmey Valle Hermosa #1 Machugal #1 LA2167 LA2580 LA1478		Cajamarca Tarapaca Tacna Apurimac Loja Loja Arequipa Amazonas Ancash Piura Amazonas Lima LA2095 LA1689 LA2184	Peru Chile Peru Peru Ecuador Ecuador Peru Peru Peru Peru Peru Peru
	LA2388 LA2746 LA3153 LA1325 LA2101 LA2863 LA3636 LA1384 LA1600 LA1687 LA2189 LA2915 LA2174 LA2963 LA1512	Cochabamba to Huambos (Chota) Asentamiento-18 Desvio Omate (Rio de Osmore) Puente Cunyac Cariamanga Macara Coayllo Quebrada Parca Las Zorras, Huarmey Valle Hermoso #2 Machugal #2 El Remanso de Olmos		Cajamarca Tarapaca Moquegua Apurimac Loja Loja Lima Lima Ancash Piura Amazonas Lambayeque LA2131 LA1924 LA2185	Peru Chile Peru Peru Ecuador Ecuador Peru Peru Peru Peru Peru Peru
	LA2548 LA2747 LA3155 LA1327 LA2103 LA2864 LA3640 LA1416 LA1601 LA1688 LA2335 LA2934 LA2204 S. peruvianum La Moyuna (Magadalena) Alta Azapa Quinistaquillas Sorocata Lansaca Sozorango Mexico City Las Delicias La Providencia Pedregal (4x) Carabayllo LA1542		Cajamarca Tarapaca Moquegua Apurimac Loja Loja Pichincha Lima Piura Lima LA2138A LA1936 LA2190	Peru Chile Peru Peru Ecuador Ecuador Mexico Ecuador Peru Peru Peru
	LA2550 LA2748 LA3355 LA1330 LA2104 LA2869 LA3781 LA1428 LA1602 LA1689 LA2340 LA2966 LA2329 LA0153 LA1545	El Tingo, Chorpampa (Rio Jequetepeque) Soledad Cacique de Ara Hacienda Francisco Pena Negra Matola-La Toma Quebrada Oscollo (Atico) La Estancilla Rio Chillon to Punchauca Castilla #1 (4x) La Molina	Cajamarca Tarapaca Tacna Apurimac Loja Loja Arequipa Manabi Lima Piura Lima LA2308 LA2102 LA2308	Peru Chile Peru Peru Ecuador Ecuador Peru Ecuador Peru Peru Peru
	LA0378 LA2553 LA2749 LA3356 LA2639B Puente Cunyac Cascas Balconcillo de San Marcos Punta Blanca W of Tacna LA2105 Jardin Botanico, Loja LA2975 Coltao LA3783 Rio Chaparra LA1466 Chongoyape LA1603 Quilca LA1690 Castilla #2 LA2345 (doubled haploid, origin unknown) LA2974 Huaca del Sol LA2409 LA0446 LA1547 LA0385 San Juan (Rio Jequetepeque) LA2555 Marical -Castilla LA2750 Mina La Despreciada LA3357 Irrigacion Magollo LA2663 Tujtohaiya LA2106 Yambra LA2976 Huangra LA3787 Alta Chaparra LA1469 El Pilar, Olmos LA1604 Horcon LA1697 Hacienda Quiroz, Santa Anita LA2346 (doubled haploid, origin unknown) LA2982 Chilca #1 LA2650 LA0752 LA1549 LA0389 Abra Gavilan LA2565 Potrero de Panacocha a Llamellin LA2751 Pachica (Rio Tarapaca) LA3358 Rio Arunta-Cono Sur LA2677 Huayapacha #1 LA2107 Los Lirios LA3794 Alta Fortaleza LA3790 Caraveli LA1470 Motupe to Desvio Olmos-Bagua LA1605 Canete -San Antonio LA1719 E of Arenillas LA2347 (doubled haploid, origin unknown) LA2983 Chilca #2 LA2864 LA1274 LA1569 LA0392 Llallan LA2566 Cullachaca LA2753 Laonzana LA3784 Rio Chaparra LA2678 Huayapacha #2 LA2108 Anganumo LA3796 Anca, Marca LA3795 Alta Fortaleza LA1471 Motupe to Jayanca LA1606 Tambo de Mora LA1720 Yautan LA2348 Trujillo LA3123 Santa Cruz: summit S. huaylasense LA1336 LA1578 LA0441 Cerro Campana LA2582 San Juan (4x) LA2754 W of Chusmisa LA3785 Terras Blancas LA2679 Huayapacha #3 LA2109 Yangana #1 LA3854 Llaguén LA3797 Anca, Marca (Rio Fortaleza) LA1472 Quebrada Topara LA1607 Canete -La Victoria LA1728 Rio San Juan LA2389 Tembladera LA3158 Los Mochis LA1364 LA1474 LA1582 LA1027 LA2583 (4x) LA2755 Banos de Chusmisa LA3786 Alta Chaparra LA2680 Puente Apurimac #1 LA2110 Yangana #2 LA3862 Purunuma LA3799 Río Pativilca LA1478 Santo Tome (Pabur) LA1608 Canete -San Luis LA1729 Rio San Juan LA2390 Chungal LA3159 Los Mochis LA1365 LA1954 LA1589 LA2917 Chullchaca LA2757 W of Chusmisa LA4106 Taltal LA2681 Puente Apurimac #2 LA2114 San Juan LA3863 Sozoranga LA3853 Mollepampa LA1514 Sayan to Churin LA1610 Asia -El Pinon LA1742 Olmos-Marquina LA2391 Chungal to Monte Grande LA3160 Los Mochis LA1982 LA2732 LA1617		Cajamarca Cajamarca Antofagasta Tacna Apurimac Loja Ancash Arequipa Lambayeque Lima Piura La Libertad LA2392 LA2173 LA2312 Cajamarca La Libertad Antofagasta Tacna Cusco Loja Ancash Arequipa Lambayeque Lima Lima Lima LA2402 LA2181 LA2411 Cajamarca Ancash Tarapaca Tacna Cusco Loja Ancash Arequipa Lambayeque Lima El Oro Lima LA2621 LA2183 LA2533 Cajamarca Ancash Tarapaca Arequipa Cusco Loja Ancash Ancash Lambayeque Ica Ancash La Libertad Galapagos Islands LA2670 LA2401 LA2560 La Libertad Cajamarca Tarapaca Arequipa Cusco Loja La Libertad Ancash Lima Lima Ica Cajamarca Sinaloa LA2675 LA2533 LA2561 Cajamarca Tarapaca Arequipa Cusco Loja Loja Ancash Piura Lima Ica Cajamarca Sinaloa LA2688 LA2852 LA2626 Ancash Tarapaca Antofagasta Cusco Loja Loja La Libertad Lima Lima Lambayeque Cajamarca Sinaloa LA2709 SolCAP LA2632	Peru Chile Peru Ecuador Peru Peru Peru Peru Peru Peru Peru Peru Peru Chile Peru Ecuador Peru Peru Peru Peru Peru Peru Peru Peru Peru Chile Peru Ecuador Peru Peru Peru Peru Ecuador Peru Peru Peru Peru Chile Peru Ecuador Peru Peru Peru Peru Peru Peru Ecuador Peru Peru Peru Chile Peru Ecuador Peru Peru Peru Peru Peru Peru Mexico Peru Peru Chile Peru Ecuador Ecuador Peru Peru Peru Peru Peru Mexico Peru Peru Peru Chile Chile Peru Ecuador Ecuador Peru Peru Peru Peru Peru Mexico
	LA4316 LA2759 LA4107 LA2695 LA2115 LA3864 LA3858 LA1520 LA1611 LA1781 LA2401 LA3161 LA2808 LA2744 LA1620	Kuntur Wasi Mamina Catarata Taltal Chihuanpampa Pucala Yangana Canta Sayan to Churin Rio Mala Bahia de Caraquez Moxeque Los Mochis		Cajamarca Tarapaca Antofagasta Cusco Loja Loja Lima Lima Lima Manabi Ancash Sinaloa LA2710 LA0166 LA2633	Peru Chile Chile Peru Ecuador Ecuador Peru Peru Peru Ecuador Peru Mexico
	LA2762 LA4108 LA3642 LA2116 LA4137 LA3900 LA1521 LA1612 LA1921 LA2412 LA3468 S. lyc. cerasiforme Quebradas de Mamina a Parca Caleta Punta Grande Ankukunka Las Juntas Barrio Delta, Cajamarca (CMV tolerant selection) El Pinon, Asia Rio Chilca Majarena Fundo Don Javier, Chilca La Molina Vieja S. pimpinellifolium LA1621		Tarapaca Antofagasta Cusco Loja Cajamarca Lima Lima Ica Lima Lima LA2783 LA0317 LA2660	Chile Chile Peru Ecuador Peru Peru Peru Peru Peru Peru
	LA2764 LA4109 LA3643 LA2119 LA4125 LA1547 LA1613 LA1923 LA2533 LA3634 LA0292 LA0373 LA1623	S. cheesmaniae (L. cheesmanii) Tarapaca Quebrada Canas Codpa Antofagasta Colcha Cusco Saraguro Loja Camina Tarapaca Chota to El Angel Carchi Santa Eusebia Lima Cabildo Ica Lomas de Latillo Lima Santa Rosa de Asia Lima LA2845 LA0373 LA2663	Chile Chile Peru Ecuador Chile Ecuador Peru Peru Peru Peru
	LA0166 LA2765 LA4117A San Pedro -Paso Jama Santa Cruz: Barranco, N of Punta Ayora Timar LA3644 Puente Tincoj LA2124 Cumbaratza LA4128 Pachica (Rio Camarones) LA1561 San Eusebio LA1614 Pampa Chumbes LA1924 Piedras Gordas LA2576 Valle de Casma LA3638 Ccatac LA1204 LA0400 LA1632		Galapagos Islands Tarapaca Antofagasta Cusco Zamora-Chinchipe Tarapaca Lima Lima Ica Ancash Lima LA2871 LA0407 LA2664	Ecuador Chile Chile Peru Ecuador Chile Peru Peru Peru Peru Peru

The word "obtained" in this sentence is quite imprecise, since it might mean "received from someone" or "obtained by breeding".

Dr H. Fukuoka, National Institute of Vegetable and Tea Science, Kusawa 360, Ano, Tsu, Mie 514-2392, Japan.

"BF-Okitsu 101" was obtained by A. Kotani, T. Kuriyama, H. Shimada-Mochizuki, S. Sakuma, and I. Suzuki (H. Fukuoka, pers. com.).

According to http://www.tofil.ph/awardee_profile.php?id=78, J Acosta won a Rockefeller scholarship to the University of Hawaii in 1958. This is also indicated in[START_REF] Acosta | Genetic analysis of bacterial wilt resistance and certain other characters in a tomato cross Lycopersicon esculentum Mill. and L. pimpinellifolium Mill[END_REF].

This hypothesis is consistent with the information of Gilbert to Laterrot(Laterrot et al., 1978) that a Philippine source had been used in Hawaii and is at the origin of "Hawaii 7996".

Peralta, I. E., D. M. Spooner, S. Knapp (2008) Taxonomy of wild tomatoes and their relatives (Solanum sect. Lycopersicoides, sect. Juglandifolia, and sect. Lycopersicon; Solanaceae). Systematic Botany Monographs 84: 1-186.

Acknowledgments

The authors are very grateful to Ph. Prior and E. Wicker (phyto-bacteriologists at CIRAD La Réunion Island, Mascarenes) for having motivated the authors to write this historical review, to F. Kaan and G. Anaïs (retirees of INRA Guadeloupe) for having provided precious complements of information, to H. Fukuoka (National Institute of Vegetables & Tea Science, Tsu, Mie, Japan) for having fully clarified the relationships between former Japanese scientists, former Japanese Institutes and tomato lines and for having translated key passages of Suzuki et al. (1964). We thank also the consortium of private companies (Vilmorin, Gautier Semences, DeRuiter Seeds, Enza Zaden, Nunhems, Rijk Zwaan) who financed (2007)(2008)(2009)(2010) research based on the present review. Last but not least, we acknowledge Ch. Olivier (librarian of INRA GAFL, Montfavet, France), M.L. Acknowledgements: This project was funded in part by grant FODECYT 54-07 to L. Mejía, by Facultad de Agronomía, Universidad de San Carlos, and by the College of Agricultural and Life Sciences, University of Wisconsin-Madison. Appreciation is expressed to Semillas Tropicales, S.A. for assistance with production of the seedlings and the field facilities.

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Conclusions:

It should be recognized that not all of the comparisons contained all of the genotypes that would have been useful for making conclusions, that viruliferous whiteflies populations would have existed at different times, that several bipartite begomoviruses as well as Tomato yellow leaf curl virus would have been present, and that weather conditions may have impacted symptom expression. In general, the following observations were considered important concerning the various introgressions: 1) Ty3a was a more effective source of resistance than Ty4. 2) Ty2 either in the homozygous or heterozygous condition was not an effective source of resistance unless it was associated with another resistance gene, such as Ty3a. 3) The heterozygous families with Ty3a were not as resistant as families that were homozygous for this introgression (Fig. 6). 4) F1 hybrids with two different introgressions were highly resistant, such as Ty3a and Ty2 (Fig. 6) or Ty3a and Ty3 (Fig. 7). 5) There was considerable variation for the DSIs among the F3 families with same genotype for an introgression, which indicated that other genes were important in conditioning resistance. This was most notable in the F3 families from the cross of Gc143-2 by Gh188-2 (Fig. 5). Others have reported that several genes are involved in highest level of begomovirus resistance expression (Anbinder et al., 2009;[START_REF] Vidavsky | Tomato breeding lines immune and tolerant to tomato yellow leaf curl virus (TYLCV) issued from Lycopersicon hirsutum[END_REF][START_REF] Zamir | Mapping and introgression of a tomato yellow leaf curl virus tolerance gene, Ty-1[END_REF] and Anbinder et al. (2009) found that there were minor QTLs for resistance associated with both the resistant and susceptible parent used in the cross for their molecular analysis of resistance. 6) For the near future it is not enough to only use molecular markers for breeding for begomovirus resistance, but field evaluations with high levels of viruliferous whiteflies will continue to be an important part of any tomato breeding program for begomovirus resistance.

Literature Cited: Abinder, I., Reuveni, M., Azari, R., Paran,I., Nahon, S., Shlomo, H., Chen, L., Lapidot, M., and Levin, I. 2009. Molecular dissection of Tomato leaf curl virus resistance in tomato line TY172 derived from Solanum peruvianum. Theor. Appl. Genet. 119:519-530. Garcia, B.E., Graham, E., Jensen, K.S., Hanson Subsequently to samples collected in December 2009, samples were collected from tomatoes with either typical begomovirus symptoms and/or purple/yellowing symptoms on younger leaves in the Department of El Progresso in March 2010. The samples were prepared for transport to the University of California-Davis using AgDia absorption strips. This is a method to capture total nucleic acids from plant tissue in which sap is prepared from the target tissue, applied to an absorbent matrix on the end of a plastic 'stick', and allowed to dry prior to transport. DNA extracts were prepared from the 12 samples and tested for the presence of Liberobacter and begomovirus infection by PCR. 'Ca. Liberibacter sp.' was detected in 4 of 12 samples, and these samples showed symptoms of stunted and distorted growth; older leaves were yellow and brittle and younger leaves were upcurled with yellowing and vein purpling. Begomovirus infection was detected in all 12 samples. Additionally, 49 tomato samples were collected from March to June, 2010 and assayed for Liberibacter sp. using PCR primers CLi.po.F/O12c [START_REF] Secor | Association of "Candidatus Liberibacter solanacearum" with Zebra Chip disease of potato established by graft and psyllid transmission, electron microscopy, and PCR[END_REF]. The expected size fragment for Liberibacter sp. was obtained from 17 samples and no fragments were obtained with the other 32 samples. Positive PCR samples were collected from the following Departments: Sacatepéquez, Guatemala and Baja Verapaz,. These PCR fragments will be sequenced for more definitive identification.

Universal PCR primers for phytoplasma [START_REF] Smart | Phytoplasm-specific PCR primers based on sequences of the 16s-23s rRNA spacer region[END_REF] were used with the DNA samples from tomato collected in Guatemala and all were negative. Because of the purple top symptoms, phytoplasma were originally suspected as being present.

In the last two years, there has been considerable effort devoted to understanding the etiology of Zebra Chip (see [START_REF] Secor | Association of "Candidatus Liberibacter solanacearum" with Zebra Chip disease of potato established by graft and psyllid transmission, electron microscopy, and PCR[END_REF] and the new Candidatus Liberibacter sp. associated with solanaceous plants (see Liefting et al., 2009aLiefting et al., , 2009b)). These efforts plus the report here indicate that this unculturable bacterium transmitted by the tomato/potato psyllid will result in a serious disease for Authors' contributions: P.S-P. designed the experiment and provided the plant material. C.L. prepared all of the specimens and measured the specimens using techniques designed in collaboration with J.SIII. C.L. drafted the Methods and Results sections, and performed the statistical analysis under the supervision of JSIII. P.S-P. drafted the Introduction and Discussion. All three authors contributed to the final editing and read and approved the manuscript before submission.