Cucurbits depicted in Byzantine mosaics from Israel, 350–600 ce

Background and Aims

Thousands of floor mosaics were produced in lands across the Roman and Byzantine empires. Some mosaics contain depictions of agricultural produce, potentially providing useful information concerning the contemporary presence and popularity of crop plants in a particular geographical region. Hundreds of floor mosaics produced in Israel during the Byzantine period have survived. The objective of the present work was to search these mosaics for Cucurbitaceae in order to obtain a more complete picture of cucurbit crop history in the eastern Mediterranean region.

Results and Conclusions

Twenty-three mosaics dating from 350–600 ce were found that had images positively identifiable as cucurbits. The morphological diversity of the cucurbit fruits in the mosaics of Israel is greater than that appearing in mosaics from any other Roman or Byzantine provincial area. The depicted fruits vary in shape from oblate to extremely long, and some are furrowed, others are striped and others lack definite markings. The cucurbit taxa depicted in the mosaics are Cucumis melo (melon), Citrullus lanatus (watermelon), Luffa aegyptiaca (sponge gourd) and Lagenaria siceraria (bottle gourd). Cucumis melo is the most frequently found taxon in the mosaics and is represented by round dessert melons and long snake melons. Fruits of at least two cultivars of snake melons and of watermelons are represented. To our knowledge, images of sponge gourds have not been found in Roman and Byzantine mosaics elsewhere. Indeed, the mosaics of Israel contain what are probably the oldest depictions of Luffa aegyptiaca in Mediterranean lands. Sponge gourds are depicted often, in 11 of the mosaics at eight localities, and the images include both mature fruits, which are useful for cleaning and washing, and immature fruits, which are edible. Only one mosaic has images positively identifiable as of bottle gourds, and these were round–pyriform and probably used as vessels.     

Autotoxic potential of cucurbit crops

Soil sickness is often observed in cucurbit crops such as Citrullus lanatus, Cucumis melo and Cucumis sativus, but not in cucurbit crops such as Cucurbita moschata, Lagenaria leucantha and Luffa cylindrica. Results showed that root aqueous extracts of Citrullus lanatus, Cucumis melo and Cucumis sativus were autotoxic, but those of Cucurbita moschata, Momordica charantia and Luffa cylindrica were less autotoxic to the radicle elongation of respective species. Plant growth of Citrullus lanatus, Cucumis melo and Cucumis sativus were greatly inhibited by autotoxic substances released from powered root tissue at a rate of 1 g per seedling. Root exudates of Citrullus lanatus, Cucumis melo and Cucumis sativus were autotoxic to radicle elongation and seedling growth of respective species. However, root exudates of Citrullus lanatus did not inhibit radicle elongation of Cucurbita ficifolia, which is commonly used as rootstock for the grafting of Citrullus lanatus, Cucumis melo and Cucumis sativus to decrease soil-borne diseases in commercial production. It seems possible to overcome autotoxicity in cucurbit crops by grafting on Cucurbita ficifolia. This revised version was published online in June 2006 with corrections to the Cover Date.     

Taxonomic significance of trichomes micromorphology in cucurbits

Studies on trichomes micromorphology using Scanning Electron Microscope (SEM) were undertaken in 23 species with one variety under 13 genera of the family Cucurbitaceae (viz., Benincasa hispida (Thunb.) Cogn., Citrullus lanatus (Thunb.) Matsum. & Nakai, Cucumis melo var. agrestis Naudin, Cucumis sativus L., Diplocyclos palmatus (L.) C. Jeffrey, Edgaria dargeelingensis C.B. Clarke, Gynostemma burmanicum King ex Chakr., Gynostemma pentaphyllum (Thunb.) Makino, Gynostemma pubescens (Gagnep.) C.Y. Wu, Hemsleya dipterygia Kuang & A.M. Lu, Lagenaria siceraria (Molina) Standl., Luffa acutangula (L.) Roxb., Luffa cylindrica M. Roem., Luffa echinata Roxb., Melothria heterophylla (Lour.) Cogn., Melothria leucocarpa (Blume) Cogn., Melothria maderspatana (L.) Cogn., Sechium edule (Jacq.) Sw., Thladiantha cordifolia (Blume) Cogn., Trichosanthes cucumerina L., T. cucumerina var. anguina (L.) Haines, Trichosanthes dioica Roxb., Trichosanthes lepiniana (Naudin) Cogn. and T. tricuspidata Lour.). The trichomes in the family Cucurbitaceae vary from unicellular to multicellular, conical to elongated, smooth to ridges, with or without flattened disk at base and cyctolithic appendages, thin to thick walled, curved at apices to blunt. Trichomes micromorphology in the family Cucurbitaceae was found significant taxonomically.     

CmMDb: A Versatile Database for Cucumis melo Microsatellite Markers and Other Horticulture Crop Research

Cucumis melo L. that belongs to Cucurbitaceae family ranks among one of the highest valued horticulture crops being cultivated across the globe. Besides its economical and medicinal importance, Cucumis melo L. is a valuable resource and model system for the evolutionary studies of cucurbit family. However, very limited numbers of molecular markers were reported for Cucumis melo L. so far that limits the pace of functional genomic research in melon and other similar horticulture crops. We developed the first whole genome based microsatellite DNA marker database of Cucumis melo L. and comprehensive web resource that aids in variety identification and physical mapping of Cucurbitaceae family. The Cucumis melo L. microsatellite database (CmMDb: http://65.181.125.102/cmmdb2/index.html) encompasses 39,072 SSR markers along with its motif repeat, motif length, motif sequence, marker ID, motif type and chromosomal locations. The database is featured with novel automated primer designing facility to meet the needs of wet lab researchers. CmMDb is a freely available web resource that facilitates the researchers to select the most appropriate markers for marker-assisted selection in melons and to improve breeding strategies.     

Medieval History of the Duda’im Melon (<Emphasis Type="Italic">Cucumis melo</Emphasis>, Cucurbitaceae)

Medieval History of the Duda’im Melon ( Cucumis melo , Cucurbitaceae). Melons, Cucumis melo, are a highly polymorphic species for fruit characteristics. The melons that are the most valued are the ones that turn sweet when ripe, including the muskmelons, cantaloupes, and casabas. Others, including the elongate adzhur, conomon, and snake melons, are consumed when immature, like cucumbers. The duda’im melons, Cucumis melo Duda’im Group, are special, as their small, spherical, thin-fleshed, insipid but beautifully maroon, dark-orange, or brown-and-yellow striped ripe fruits are valued for ornament and especially for their lush fragrance. The distinctive properties of duda’im melons are matched with special names given to them in several languages and geographical areas, which have made possible tracing of the history of these melons to mid-9^th century Persia. From that region, duda’im melons diffused westward, likely facilitated by Islamic conquests, reaching North Africa and Andalusia in the 10^th century.     

A short history of Lagenaria siceraria (bottle gourd) in the Roman provinces: morphotypes and archaeogenetics

Lagenaria siceraria (Molina) Standl. (bottle gourd) is pantropic and displays large variation in fruit and seed shape. Two subspecies are currently recognized: the African L. siceraria ssp. siceraria and the Asian L. siceraria ssp. asiatica. The Asian type of bottle gourd belongs to the earliest domesticated plants in the Americas. In Europe, bottle gourd only appears with some frequency from the Roman period onwards. The paper is the study of ancient DNA (aDNA) and seed morphology of one almost complete bottle gourd fruit from the Roman site of Oedenburg/Biesheim–Kunheim, France (1st century a.d.), and from individual seed finds from the Roman vicus of Petinesca–Vorderberg, Switzerland (3rd century a.d.), both recovered from waterlogged layers. Width and length measurements of seeds show large variation. Based on the index of width to length, seeds from both sites differ significantly (p?<?0.0001 Mann–Whitney) suggesting that there were different variants present north of the Alps. Genetically, the bottle gourd fruit from Roman Oedenburg/Biesheim–Kunheim and one commercial cultivar L. siceraria cv. ‘Herkuleskeule’ are of Asian origin as identified by three Asian and African specific chloroplast markers. These results support an early and long-lasting presence of the Asian type of domestic bottle gourd in Europe. No chloroplast markers were found in the seeds from Petinesca–Vorderberg. However preserved nuclear high copy 5.8S rDNA fragments correctly matched to Cucurbitaceae, further supporting the evidence for preservation of DNA in waterlogged plant remains.     

Sequencing of 6.7 Mb of the melon genome using a BAC pooling strategy

Background  

Cucumis melo (melon) belongs to the Cucurbitaceae family, whose economic importance among horticulture crops is second only to Solanaceae. Melon has a high intra-specific genetic variation, morphologic diversity and a small genome size (454 Mb), which make it suitable for a great variety of molecular and genetic studies. A number of genetic and genomic resources have already been developed, such as several genetic maps, BAC genomic libraries, a BAC-based physical map and EST collections. Sequence information would be invaluable to complete the picture of the melon genomic landscape, furthering our understanding of this species' evolution from its relatives and providing an important genetic tool. However, to this day there is little sequence data available, only a few melon genes and genomic regions are deposited in public databases. The development of massively parallel sequencing methods allows envisaging new strategies to obtain long fragments of genomic sequence at higher speed and lower cost than previous Sanger-based methods.     

Overview of the origins and history of the five major cucurbit crops: issues for ancient DNA analysis of archaeological specimens

The gourd family, Cucurbitaceae, contains five vegetable crops of worldwide importance, the pumpkins and squash (Cucurbita spp.), watermelons (Citrullus lanatus), melons (Cucumis melo) and cucumbers (Cucumis sativus). Here is presented a synopsis of the origin and history of these cucurbit crops. Historical records of the use of cucurbits by people take the form of archaeobotanical finds, iconography and literature. The weight of the evidence indicates that Cucurbita spp. were first cultivated in the Americas at least 10,000 years ago, and that by 1492 ce a number of cultivar-groups of pumpkins and squash had been developed by indigenous American peoples. Watermelons were cultivated in northeastern Africa at least 4,000 years ago, first probably as a source of fresh water. Melons and cucumbers are native to Asia, probably initially cultivated for the use of the young fruits as vegetables. Melons spread to eastern Africa at an early date, but cucumbers are probably a more recent domesticate and spread westwards later, reaching Europe in early medieval times. Sequencing of cucurbit plant genomes and advances in ancient DNA research offer much promise for obtaining an improved assessment of cucurbit crop origins, specifically the genetic constitution and geographical home of ancestral source populations. Next-generation genomic sequencing, if applied to an appropriate array of archaeological cucurbit remains and modern germplasm, could contribute much to the understanding of the history and evolution under domestication of cucurbit crops.     

Bipolaris multiformis (Jooste) Alcorn Pathogenic on Leaves of Cucumis sativus in Italy

A destructive disease was observed in August 1981 in a field of Cucumis sativus cv. Radar. Sincc then sporadic spots on leaves of the same cultivar were recorded in 1985, 1988 and 1993. The causal agent was identified in Bipolaris multiformis (Jooste) Alcorn. The fungus produces frequently irregular (mostly Y-shaped) conidia in culture but rarely on leaves. The pathogen germinates bipolarly in distilled water at 20–25°C (95 and 90% respectively), and on leaves it produces appressoria;, entry points are either stomata or the intact surface. Pathogenicity tests carried out on various cucurbitaceous plants showed symptom intensity decreasing with the following order: C. sativus cv. Radar and Eeballium elaterium > Brionia cretica subsp. dioica > Cueurbita pepo L. var. pommiformis and C. pepo var. patissonina f. radiata > C. moscata and Sechium edule. The other two tested cultivars of Cucumis sativus (Mezzolungo Marketer and Corto per sottaceti) and C. pepo cv. Verde scuro di Milano showed sterile, very small necrotic dot. Control plants, Citrullus lanatus cv. Sugar baby, Cucumbis melo and Lagenaria siceraria showed no symptoms.     

24β-Ethyl-31-norlanosta-8,25(27)-dien-3β-ol and 24β-ethyl-25(27)-dehydrolophenol in seeds of three cucurbitaceae species

The structures of two 4α-methylsterols is isolated from Cucumis sativus(Cucurbitaceae) seeds were determined based mainly on their ¹³CNMR spectra as 24β-ethyl-31-norlanosta-8,25(27)-dien-3β-ol and 24β-ethyl-25(27)- dehydrolophenol, respectively, of which the former is a new sterol from natural sources. These two 4α-methylsterols were identified in the seeds of two other Cucurbitaceae species, Lagenaria leucantha var. Gourda and Citrullus battich. The probable biogenetic significance of the two 4α-methylsterols is discussed. Other 4α-methylsterols identified in the seeds of the three Cucurbitaceae species were obtusifoliol, cycloeucalenol and gramisterol.     

A tobacco ringspot virus-based vector system for gene and microRNA function studies in cucurbits

Cucurbits are economically important crops worldwide. The genomic data of many cucurbits are now available. However, functional analyses of cucurbit genes and noncoding RNAs have been impeded because genetic transformation is difficult for many cucurbitaceous plants. Here, we developed a set of tobacco ringspot virus (TRSV)-based vectors for gene and microRNA (miRNA) function studies in cucurbits. A TRSV-based expression vector could simultaneously express GREEN FLUORESCENT PROTEIN (GFP) and heterologous viral suppressors of RNA silencing in TRSV-infected plants, while a TRSV-based gene silencing vector could knock down endogenous genes exemplified by PHYTOENE DESATURASE (PDS) in Cucumis melo, Citrullus lanatus, Cucumis sativus, and Nicotiana benthamiana plants. We also developed a TRSV-based miRNA silencing vector to dissect the functions of endogenous miRNAs. Four representative miRNAs, namely, miR159, miR166, miR172, and miR319, from different cucurbits were inserted into the TRSV vector using a short tandem target mimic strategy and induced characteristic phenotypes in TRSV-miRNA-infected plants. This TRSV-based vector system will facilitate functional genomic studies in cucurbits.     

Long-read genome assembly and genetic architecture of fruit shape in the bottle gourd

The bottle gourd (Lagenaria siceraria, Cucurbitaceae) is an important horticultural crop exhibiting tremendous diversity in fruit shape. The genetic architecture of fruit shape variation in this species remains unknown. We assembled a long-read-based, high-quality reference genome (ZAAS_Lsic_2.0) with a contig N50 value over 390-fold greater than the existing reference genomes. We then focused on dissection of fruit shape using a one-step geometric morphometrics-based functional mapping approach. We identified 11 quantitative trait loci (QTLs) responsible for fruit shape (fsQTLs), reconstructed their visible effects and revealed syntenic relationships of bottle gourd fsQTLs with 12 fsQTLs previously reported in cucumber, melon or watermelon. Homologs of several well-known and newly identified fruit shape genes, including SUN, OFP, AP2 and auxin transporters, were comapped with bottle gourd QTLs.     

Length polymorphism and homologies of microsatellites in several Cucurbitaceae species

The objectives of this research were to assess (1) the degree of Simple Sequence Repeats (SSR) DNA length polymorphism in melon (Cucumis melo L.) and other species within the Cucurbitaceae family and (2) the possibility of utilizing SSRs flanking primers from single species to other genera or species of Cucurbitaceae. Five melon (CT/GA) _n SSRs were isolated from a genomic library. Two cucumber (Cucumis sativus L.) SSRs were detected through a search of DNA sequence databases, one contained a (CT)₈ repeat, the other a (AT)₁₃ repeat. The seven SSRs were used to test a diverse sample of Cucurbitaceae, including 8 melon, 11 cucumber, 5 squash, 1 pumpkin, and 3 watermelon genotypes. Five of the seven SSRs detected length polymorphism among the 8 melon genotypes. PCR amplification revealed between three and five length variants (alleles) for each SSR locus, with gene diversity values ranging from 0.53 to 0.75. Codominant segregation of the alleles among F₂ progeny was demonstrated for each of the five SSR loci. Four of the seven SSRs detected polymorphism among the 11 cucumber genotypes, with gene diversity values ranging between 0.18 and 0.64. Primers specific to SSRs of C. melo and C. sativus also amplified DNA extracted from genotypes belonging to other genera of the Cucurbitaceae family.     

Genetic diversity of cucumber estimated by morpho-physiological and EST-SSR markers

In the present study, genetic variation among 40 cucumber genotypes was analyzed by means of morpho-physiological traits and 21 EST-SSR markers. Diversity was observed for morpho-physiological characters like days to 50% female flowering (37–46.9, number of fruits/plant (1.33–5.80), average fruit weight (41–333), vine length (36–364), relative water content (58.5–92.7), electrolyte leakage (15.9–37.1), photosynthetic efficiency (0.40–0.75) and chlorophyll concentration index (11.1–28.6). The pair wise Jaccard similarity coefficient ranged from 0.00 to 0.27 for quantitative traits and 0.24 to 0.96 for EST-SSR markers indicating that the accessions represent genetically diverse populations. With twenty-one EST-SSR markers, polymorphism revealed among 40 cucumber genotypes, number of alleles varied 2–6 with an average 3.05. Polymorphism information content varied from 0.002 to 0.989 (mean = 0.308). The number of effective allele (Ne), expected heterozygosity (He) and unbiased expected heterozygosity (uHe) of these EST-SSRs were 1.079–1.753, 0.074–0.428 and 0.074–0.434, respectively. Same 21 EST-SSR markers transferability checked in four other Cucumis species: snapmelon (Cucumis melo var. momordica), muskmelon (Cucumis melo L.), pickling melon (Cucumis melo var. conomon) and wild muskmelon (Cucumis melo var. agrestis) with frequency of 61.9, 95.2, 76.2, and 76.2%, respectively. Present study provides useful information on variability, which can assist geneticists with desirable traits for cucumber germplasm utilization. Observed physiological parameters may assists in selection of genotype for abiotic stress tolerance also, EST-SSR markers may be useful for genetic studies in related species.     

Transferability of Cucurbita SSR markers for genetic diversity assessment of Turkish bottle gourd (Lagenaria siceraria) genetic resources

The genetic diversity present in crop landraces represents a valuable genetic resource for breeding and genetic studies. Bottle gourd (Lagenaria siceraria) landraces in Turkey are highly genetically diverse. However, the limited genomic resources available for this crop hinder the molecular characterization of Turkish bottle gourd germplasm for its adequate conservation and management. Therefore, we evaluated the efficacy of 40 SSR markers from major cucurbit crops (Cucurbita pepo L. and Cucurbita moschata L.) in 30 bottle gourd landraces, together with 16 SRAP primer combinations. In addition, we compared the genetic relationship between bottle gourd and 31 other cucurbit accessions (11 Cucurbita maxima, 3 C. moschata, 5 C. pepo subsp. ovifera, 10 C. pepo and 2 Luffa cylindrica). Twenty-seven Cucurbita SSR markers showed transferability to bottle gourd. SSR markers amplified 59 alleles, in bottle gourd genome with an average of 1.64 alleles per locus. Together, SSR and SRAP markers amplified 453 fragments across the 61 accessions, and clearly discriminated L. siceraria and L. cylindrica from the other cucurbit species. Genetic diversity analysis separated edible cucurbit from ornamentals, while population structure analysis classified L. siceraria in two subpopulations defined by fruit shape, rather than geographical origin. The results indicated that the genomic resources available for Cucurbita species are valuable to study and preserve the genetic diversity of bottle gourd in Turkey.     

Entomofaune associée à la culture de cucurbites oléagineuses africaines (Lagenaria siceraria Molina (Standl. 1930) et Citrullus lanatus Thumb (Matsum & Nakai 1916)) et impact des ravageurs sur la production

Les observations faites sur deux cucurbitacées africaines, Lagenaria siceraria Molina (Standl.) et Citrullus lanatus Thumb (Matsum & Nakai) ont permis d’inventorier l’entomofaune (utiles et ravageurs) associée à la pratique culturale. L’évaluation des dégâts occasionnés par les ravageurs a mis en évidence leur impact sur la production. Suivant un schéma expérimental, des parcelles de 49 m² traitées au CYPERCAL 50 EC (50 g/l de Cypermétrine) et au FURADAN 5G (50 g/kg de Carbofuran) et non traitées ont été disposées en blocs aléatoires. Les observations ont porté sur l’identification et le dénombrement des insectes, sur le pourcentage de plants attaqués ainsi que sur la production de baies mûres et de graines. Au total, 71 espèces ont été identifiées, réparties en 41 familles et 10 ordres (Coléoptères, Hyménoptères, Diptères, Lépidoptères, Orthoptères, Homoptères, Hétéroptères, Dictyoptères, Odonates et Dermaptères). Les principaux ravageurs appartiennent à la famille des Chrysomelidae (Lamprocopa occidentalis, Aulacophora foveicolis, Aulacophora africana, Asbecesta cyanipennis, Ootheca mutabilis et Lilioceris livida), des Coccinellidae (Henosepilachna elaterii et Henosepilachna reticulata) et des Meloidae (Mylabris holosericea). Plus de 50 % de plants sains ont été recensés dans les parcelles traitées contre 0 à 33 % dans les parcelles témoins. Les nombres moyens des baies obtenues dans les parcelles traitées (37,16 ± 2,99 pour L. siceraria et 42,33 ± 2,58 pour C. lanatus) sont significativement (P < 0,001) plus importants que ceux observés dans les parcelles témoins (29,33 ± 4,67 pour L. siceraria et 31,33 ± 3,5 pour C. lanatus). Les rendements moyens en graines dans les parcelles traitées (222,39 ± 15,78 kg/ha pour L. siceraria et 127,4 ± 8,16 kg/ha pour C. lanatus) sont significativement (P < 0,001) supérieurs à ceux observés dans les parcelles témoins (144,4 ± 14,52 kg/ha pour L. siceraria et 88,18 ± 8,09 kg/ha pour C. lanatus). Les traitements insecticides, en réduisant les populations de ravageurs et les dégâts sur les plants, ont entrainé une augmentation du rendement à l’hectare de plus de 40 %.     

Cucurbit Grafting

Due to limited availability of arable land and high market demand for off-season vegetables, cucurbits (plants in the family Cucurbitaceae) are continuously cultivated under unfavorable conditions in some countries. These conditions include environments that are too cold, wet, or dry, or are cool low-light winter greenhouses. Successive cropping can increase salinity, the incidence of cucurbit pests, and soilborne diseases like fusarium wilt caused by Fusarium spp. These conditions cause various physiological and pathological disorders leading to severe crop loss. Chemical pest control is expensive, not always effective, and can harm the environment. Grafting can overcome many of these problems. In fact, in many parts of the world, grafting is a routine technique in continuous cropping systems. It was first commonly used in Japan during the late 1920s by grafting watermelon [Citrullus lanatus (Thunb.) Matsum. and Nakai] onto pumpkin [Cucurbita moschata Duchesne ex. Poir] rootstocks. Soon after, watermelons were grafted onto bottle gourd [Lagenaria siceraria (Molina) Standl.] rootstocks. This practice helped control declining yield due to soilborne diseases. China produces more than half the world's watermelons and cucumbers (Cucumis sativus L.), and approximately 20% of these are grafted. Use of rootstocks can enhance plant vigor through vigorous attainment of soil nutrients, avoidance of soil pathogens and tolerance of low soil temperatures, salinity, and wet-soil conditions. The type of rootstock affects cucurbit plant growth, yield, and fruit quality. Cucurbit grafting is rare in the United States, but with continued loss of quality disease-free farmland along with the phase-out of methyl bromide, the U.S. cucurbit industry sees grafting as an attractive option. Some seed companies now offer watermelon transplants grafted onto squash or bottle gourd rootstocks, and some transplant facilities offer grafting services. There have been thorough analyses of cucurbit grafting in other countries, but the literature in English is limited. This review summarizes the state of the cucurbit grafting industry on a global level, translating work published in many languages.