Flower morphology and floral sequence in Artemisia annua (Asteraceae)1.

UNLABELLED: • PREMISE OF THE STUDY: Artemisia annua produces phytochemicals possessing antimalarial, antitumor, anti-inflammatory, and anthelmintic activities. The main active ingredient, artemisinin, is extremely effective against malaria. Breeding to develop cultivars producing high levels of artemisinin can help meet worldwide demand for artemisinin and its derivatives. However, fundamental reproductive processes, such as the sequence of flowering and fertility, are not well understood and impair breeding and seed propagation programs.• METHODS: Capitulum structure and floral sequence were studied using light and scanning electron microscopy to describe inflorescence architecture, floret opening, and seed set.• KEY RESULTS: Florets are minute and born in capitula containing pistillate ray florets and hermaphroditic disk florets. Ray florets have elongated stigmatic arms that extend prior to disk floret opening. Disk florets exhibit protandry. During the staminate phase, pollen is released within a staminate tube and actively presented with projections at the tip of stigmas as the pistil elongates. During the pistillate phase, stigmatic arms bifurcate and reflex. Stigmas are of the dry type and stain positively for polysaccharides, lipids, and an intact cuticle. Floret numbers vary with genotype, and capitula are predominantly composed of disk florets. Both ray and disk florets produce filled seed.• CONCLUSIONS: Gynomonoecy, early opening of ray florets, and dichogamy of disk florets promote outcrossing in A. annua For breeding and seed development, flowering in genotypes can be synchronized under short days according to the floral developmental stages defined. Floret number and percentage seed fill vary with genotype and may be a beneficial selection criterion.

Postharvest physiology and technology of loquat (Eriobotrya japonica Lindl.) fruit.

Loquat (Eriobotrya japonica Lindl.) is a subtropical evergreen tree whose fruit is consumed both fresh and processed. Loquat fruit is a good source of minerals and carotenoids, while the kernel is rich in protein and carbohydrates. It has been considered a non-climacteric fruit, but there is evidence that some cultivars have a ripening pattern similar to that of climacteric fruits. The fruit has a short postharvest life at ambient temperatures and is susceptible to physical and mechanical damage, loss of moisture and nutrients, and decay. Low-temperature storage extends the shelf life of loquat fruit, but some cultivars are severely affected by chilling injury and flesh browning during cold storage. Purple spot, browning and leatheriness are major postharvest disorders. The shelf life of loquat can be extended by modified or controlled atmosphere storage as well as by postharvest treatment with 1-methyl cyclopropene or methyl jasmonate.

Medieval iconography of watermelons in Mediterranean Europe.

BACKGROUND AND AIMS: The watermelon, Citrullus lanatus (Cucurbitaceae), is an important fruit vegetable in the warmer regions of the world. Watermelons were illustrated in Mediterranean Antiquity, but not as frequently as some other cucurbits. Little is known concerning the watermelons of Mediterranean Europe during medieval times. With the objective of obtaining an improved understanding of watermelon history and diversity in this region, medieval drawings purportedly of watermelons were collected, examined and compared for originality, detail and accuracy. FINDINGS: The oldest manuscript found that contains an accurate, informative image of watermelon is the Tractatus de herbis, British Library ms. Egerton 747, which was produced in southern Italy, around the year 1300. A dozen more original illustrations were found, most of them from Italy, produced during the ensuing two centuries that can be positively identified as watermelon. In most herbal-type manuscripts, the foliage is depicted realistically, the plants shown as having long internodes, alternate leaves with pinnatifid leaf laminae, and the fruits are small, round and striped. The manuscript that contains the most detailed and accurate image of watermelon is the Carrara Herbal, British Library ms. Egerton 2020. In the agriculture-based manuscripts, the foliage, if depicted, is not accurate, but variation in the size, shape and coloration of the fruits is evident. Both red-flesh and white-flesh watermelons are illustrated, corresponding to the typical sweet dessert watermelons so common today and the insipid citron watermelons, respectively. The variation in watermelon fruit size, shape and coloration depicted in the illustrations indicates that at least six cultivars of watermelon are represented, three of which probably had red, sweet flesh and three of which appear to have been citrons. Evidently, citron watermelons were more common in Mediterranean Europe in the past than they are today.

Occidental diffusion of cucumber (Cucumis sativus) 500-1300 CE: two routes to Europe.

BACKGROUND: The cucumber, Cucumis sativus, is one of the most widely consumed fruit vegetables the world over. The history of its dispersal to the Occident from its centre of origin, the Indian subcontinent, has been incorrectly understood for some time, due to the confusion of cucumbers with vegetable melons. Iconographic and literary evidence has shown that cucumber was absent in Roman times, up to 500 CE, but present in Europe by late medieval times, 1300. The objective of the present investigation was to determine more accurately when the cucumber arrived in Europe and by what route. FINDINGS AND CONCLUSIONS: The evidence for the movement of C. sativus westward is entirely lexicographical until the 10th century. Syriac, Persian and Byzantine Greek sources suggest the presence of cucumbers, to the east and north-east of the Mediterranean Sea (modern Iran, Iraq and Turkey), by the 6th or 7th century. Arabic medical writings suggest the presence of cucumbers in Spain as early as the mid-9th century and in Tunisia by the early 10th century. Descriptive evidence in Arabic establishes the presence of cucumbers in Andalusia by the second half of the 10th century. Latin translations from Arabic sources indicate the presence of cucumbers in southern Italy by the second half of the 11th century. These writings, together with lexicographical discrepancies in names of cucurbits in late medieval Latin writings, suggest that cucumber was introduced to Europe by two independent diffusions. One diffusion appears to have been overland from Persia into eastern and northern Europe and preceded the Islamic conquests. The other, subsequent diffusion into western and southern Europe, was probably by a mostly maritime route from Persia or the Indian subcontinent into Andalusia.

Iconography of Beans and Related Legumes Following the Columbian Exchange.

Common bean (Phaseolus vulgaris L.), maize, and squash were described by explorers as early as 1492. The illustration of common bean recognized as the first in Europe is in Fuchs' Di Historias Stirpium, published in 1542 and a half-century after beans were observed in the Caribbean. Besides herbals and herbarium specimens, the sources of information on the introduction of New World crops are paintings and illustrations. Two early sources of images of maize and squash are the Grandes Heures d'Anne de Bretagne and the Loggia di Amore e Psiche in the Villa Farnesina, Rome. The former was illustrated between 1507 and 1508 and has an image identified as the common bean. The Villa Farnesina Loggia was decorated in 1515-1518, with festoons containing three instances of bean pods. Our first objective was to evaluate these images to determine whether they represented depictions of common bean earlier than the illustration by Fuchs. Neither image appears to be a common bean based on a combination of botanical characters and size. Folio 194 of the Grandes Heures d'Anne de Bretagne is most likely a Vigna species in the Ceratotropis subgenus. In the Loggia, one set of pods appears to be a species in the Mimosoideae subfamily and the second and third sets of pods most closely resemble Canavalia gladiata. Neither image likely represents common beans and are probably Old-World species. Secondly, illustrations of common beans from ten early herbals were analyzed for traits that are characteristic of the centers of domestication and races of common beans. Our objective was to characterize the diversity observed among herbals and determine whether beans from both centers of domestication were present. We potentially identified both Middle American, race Mesoamerica and Andean, race Nueva Granada types. We posit that both Middle American and Andean types were in the Caribbean at the time of the Columbian exchange and that beans from both centers were informally introduced into Europe early on. This review of 16th-century manuscripts and illustrations has provided some answers to the questions of what and when common beans reached Europe and provide new hypotheses for researchers studying the origins, diversity, and distribution of this crop.

Medieval herbal iconography and lexicography of Cucumis (cucumber and melon, Cucurbitaceae) in the Occident, 1300-1458.

BACKGROUND: The genus Cucumis contains two species of important vegetable crops, C. sativus, cucumber, and C. melo, melon. Melon has iconographical and textual records from lands of the Mediterranean Basin dating back to antiquity, but cucumber does not. The goal of this study was to obtain an improved understanding of the history of these crops in the Occident. Medieval images purportedly of Cucumis were examined, their specific identity was determined and they were compared for originality, accuracy and the lexicography of their captions. FINDINGS: The manuscripts having accurate, informative images are derived from Italy and France and were produced between 1300 and 1458. All have an illustration of cucumber but not all contain an image of melon. The cucumber fruits are green, unevenly cylindrical with an approx. 2:1 length-to-width ratio. Most of the images show the cucumbers marked by sparsely distributed, large dark dots, but images from northern France show them as having densely distributed, small black dots. The different size, colour and distribution reflect the different surface wartiness and spininess of modern American and French pickling cucumbers. The melon fruits are green, oval to serpentine, closely resembling the chate and snake vegetable melons, but not sweet melons. In nearly all manuscripts of Italian provenance, the cucumber image is labelled with the Latin caption citruli, or similar, plural diminuitive of citrus (citron, Citrus medica). However, in manuscripts of French provenance, the cucumber image is labelled cucumeres, which is derived from the classical Latin epithet cucumis for snake melon. The absence of melon in some manuscripts and the expropriation of the Latin cucumis/cucumer indicate replacement of vegetable melons by cucumbers during the medieval period in Europe. One image, from British Library ms. Sloane 4016, has a caption that allows tracing of the word 'gherkin' back to languages of the geographical nativity of C. sativus, the Indian subcontinent.

Evolution of Root Morphology in Table Beet: Historical and Iconographic.

The Beta vulgaris complex includes sugar beet, mangel wurzel, Swiss chard, fodder beet, and table beet. Mangel wurzel and fodder beet are considered to be the same general crop type, with the former possessing lower dry matter content (<13%) than the latter. Mangel is likely derived from crosses between table beet and chard, while fodder beet may have a more recent origin, arising from crosses between mangel and sugarbeet. The table beet was derived from the wild sea beet, B. vulgaris (L.) subsp. maritima (L.) Arcang, with small non-spherical roots. Table beet is presently a popular vegetable cultivated for its pigmented roots, typically red but also yellow and other colors. Wild forms were consumed in antiquity mainly for their leaves with roots used medicinally. Beet is referred to in the Septuagint, a Greek translation of the first five books of the Hebrew bible, made in Ptolomeic Egypt in the third century BCE. A beet identified as Beta maritima is included in De Material Medicus of Pedanius Dioscorides written in the first century CE, and the first illustrated version of 512, known as the Juliana Anicia Codex, includes an image with non-spherical root. Beet is mentioned in several tractates of the Talmud, a sixth century collection of history and civil law written in Babylonia. Beta maritima possesses supernumerary root cambia, which facilitated selection of swollen rooted forms. The first colored illustration of swollen rooted table beet, B. vulgaris, can be found in the 1515-1517 frescos of Raphael Sanzio and Giovanni Martina da Udine in the Villa Farnesina in Rome. Swollen roots in Roman beet are illustrated and described in the 1587 French herbal Historia Generalis Plantarum of Jacques Dalechamps. Conically shaped beet roots are found in the market painting of Franz Snijders in the 17th century. Various spherical forms of beet root are found in the work of American painter James Peale in 1826. A complete array of beet root types is found in the Benary catalog of 1876. Modern, spherical beet roots were depicted in 1936 by the Russian painter Zinaida Serebriankov, 1936. Artistic and historical representations of table beet suggest that swollen rooted forms have existed during the past five centuries, but conically shaped roots were gradually replaced by spherically shaped roots during this period.

The Cucurbitaceae and Solanaceae illustrated in medieval manuscripts known as the Tacuinum Sanitatis.

BACKGROUND AND AIMS: Beginning in the last two decades of the 14th century, richly illuminated versions of the Tacuinum Sanitatis, the Latin translation of an 11th-century Arabic manuscript known as Taqwim al-Sihha bi al-Ashab al-Sitta, were produced in northern Italy. These illustrated manuscripts provide a window on late medieval life in that region by containing some 200 full-page illustrations, many of which vividly depict the harvest of vegetables, fruits, flowers, grains, aromatics and medicinal plants. Our objective was to search for and identify the images of taxa of Cucurbitaceae and Solanaceae. METHODS: We have located all reported illustrated Tacuinum Sanitatis and similar or related manuscripts, searched through printed or electronic reproductions of them, categorized six of them that display full-page illustrations as archetypic, and established the identity of the Cucurbitaceae and Solanaceae appearing in these six manuscripts. KEY RESULTS AND CONCLUSIONS: Of the Cucurbitaceae, Cucumis sativus (short-fruited cucumbers), Cucumis melo (including round as well as elongate melons), Citrullus lanatus (both sweet watermelons and citrons), and Lagenaria siceraria (including bottle-shaped as well as long gourds), are illustrated. Of the Solanaceae, Solanum melongena (egg-shaped purple aubergines) and Mandragora sp. (mandrake) are illustrated. These depictions include some of the earliest known images of cucumber, casaba melon (Cucumis melo Inodorous Group) and aubergine, each of which closely resembles an extant cultivar-group or market type. Overall, the botanically most accurate images are in the version of the Tacuinum located in the Osterreichische Nationalbibliothek, Vienna, cod. ser. n. 2644. Similarities and differences in botanical accuracy among the images of Cucurbitaceae and Solanaceae in the six archetypal Tacuinum manuscripts suggest to us that another illustrated Tacuinum, now lost, may have antedated and served as a model or inspiration for the six surviving archetypic manuscripts.

Selection and Clonal Propagation of High Artemisinin Genotypes of Artemisia annua.

Artemisinin, produced in the glandular trichomes of Artemisia annua L. is a vital antimalarial drug effective against Plasmodium falciparum resistant to quinine-derived medicines. Although work has progressed on the semi-synthetic production of artemisinin, field production of A. annua remains the principal commercial source of the compound. Crop production of artemisia must be increased to meet the growing worldwide demand for artemisinin combination therapies (ACTs) to treat malaria. Grower artemisinin yields rely on plants generated from seeds from open-pollinated parents. Although selection has considerably increased plant artemisinin concentration in the past 15 years, seed-generated plants have highly variable artemisinin content that lowers artemisinin yield per hectare. Breeding efforts to produce improved F1 hybrids have been hampered by the inability to produce inbred lines due to self-incompatibility. An approach combining conventional hybridization and selection with clonal propagation of superior genotypes is proposed as a means to enhance crop yield and artemisinin production. Typical seed-propagated artemisia plants produce less than 1% (dry weight) artemisinin with yields below 25 kg/ha. Genotypes were identified producing high artemisinin levels of over 2% and possessing improved agronomic characteristics such as high leaf area and shoot biomass production. Field studies of clonally-propagated high-artemisinin plants verified enhanced plant uniformity and an estimated gross primary productivity of up to 70 kg/ha artemisinin, with a crop density of one plant m-2. Tissue culture and cutting protocols for the mass clonal propagation of A. annua were developed for shoot regeneration, rooting, acclimatization, and field cultivation. Proof of concept studies showed that both tissue culture-regenerated plants and rooted cutting performed better than plants derived from seed in terms of uniformity, yield, and consistently high artemisinin content. Use of this technology to produce plants with homogeneously-high artemisinin can help farmers markedly increase the artemisinin yield per cultivated area. This would lead to increased profit to farmers and decreased prices of ACT.

The cucurbits of mediterranean antiquity: identification of taxa from ancient images and descriptions.

BACKGROUND: A critical analysis was made of cucurbit descriptions in Dioscorides' De Materia Medica, Columella's De Re Rustica and Pliny's Historia Naturalis, works on medicine, agriculture and natural science of the 1st century ce, as well as the Mishna and Tosefta, compilations of rabbinic law derived from the same time period together with cucurbit images dating from antiquity including paintings, mosaics and sculpture. The goal was to identify taxonomically the Mediterranean cucurbits at the time of the Roman Empire. FINDINGS: By ancient times, long-fruited forms of Cucumis melo (melon) and Lagenaria siceraria (bottle gourd) were selected, cultivated and used as vegetables around the Mediterranean and, in addition, bottle-shaped fruits of L. siceraria were employed as vessels. Citrullus lanatus (watermelons) and round-fruited forms of Cucumis melo (melons) were also consumed, but less commonly. A number of cucurbit species, including Bryonia alba, B. dioica, Citrullus colocynthis and Ecballium elaterium, were employed for medicinal purposes. No unequivocal evidence was found to suggest the presence of Cucumis sativus (cucumber) in the Mediterranean area during this era. The cucumis of Columella and Pliny was not cucumber, as commonly translated, but Cucumis melo subsp. melo Flexuosus Group (snake melon or vegetable melon).

The cucurbit images (1515-1518) of the Villa Farnesina, Rome.

BACKGROUND: The gorgeous frescoes organized by the master Renaissance painter Raphael Sanzio (1483-1520) and illustrating the heavenly adventures of Cupid and Psyche were painted between 1515 and 1518 to decorate the Roman villa (now known as the Villa Farnesina) of the wealthy Sienese banker Agostino Chigi (1466-1520). Surrounding these paintings are festoons of fruits, vegetables and flowers painted by Giovanni Martini da Udine (1487-1564), which include over 170 species of plants. A deconstruction and collation of the cucurbit images in the festoons makes it possible to evaluate the genetic diversity of cucurbits in Renaissance Italy 500 years ago. FINDINGS: The festoons contain six species of Old World cucurbits, Citrullus lanatus (watermelon), Cucumis melo (melon), Cucumis sativus (cucumber), Ecballium elaterium (squirting cucumber), Lagenaria siceraria (bottle gourd) and Momordica balsamina (balsam apple), and two or three species of New World cucurbits, Cucurbita maxima, C. pepo and, perhaps, C. moschata (pumpkin, squash, gourd). The images of C. maxima are the first illustrations of this species in Europe.

First known image of Cucurbita in Europe, 1503-1508.

BACKGROUND: The genus Cucurbita (pumpkin, squash, gourd) is native to the Americas and diffused to other continents subsequent to the European contact in 1492. For many years, the earliest images of this genus in Europe that were known to cucurbit specialists were the two illustrations of C. pepo pumpkins that were published in Fuchs' De Historia Stirpium, 1542. Images of fruits of two Cucurbita species, drawn between 1515 and 1518, were recently discovered in the Villa Farnesina in Rome. FINDINGS: An even earlier image of Cucurbita exists in the prayer book, Grandes Heures d'Anne de Bretagne, illustrated by Jean Bourdichon in Touraine, France, between 1503 and 1508. This image, which shows a living branch bearing flowers and fruits, had not been examined and analysed by cucurbit specialists until now. The image is identified as depicting Cucurbita pepo subsp. texana. Unlike some of the fruits of Cucurbita depicted in the Villa Farnesina a decade later, this image does not depict an esculent and does not constitute evidence of early European contact with New World agriculture. Based on the descriptive, ecological and geographical accounts of C. pepo subsp. texana in the wild, the idea is considered that the image was based on an offspring of a plant found growing along the Gulf Coast of what is now the United States.