PHYTOPLASMA DISEASES OF MEDICINAL AND AROMATIC PLANTS

Offered review

PHYTOPLASMA DISEASES OF MEDICINAL AND AROMATIC PLANTS

C. Marcone1, M.G. Bellardi2 and A. Bertaccini2

1 Department of Pharmacy, University of Salerno, Via Giovanni Paolo II, 132, I-84084 Fisciano (Salerno), Italy2 Department of Agricultural Science, Alma Mater Studiorum - University of Bologna, Viale G. Fanin, 42, I-40127 Bologna, Italy

SUMMARY

Medicinal and aromatic plants include a broad array of wild and cultivated plants which contain many biolog-ically-active compounds, known as phytochemicals, that are of great interest for their ability to promote human and animal health. The present review provides a litera-ture overview of phytoplasma diseases affecting medicinal and aromatic plants, with an emphasis on phytoplasma taxa associated. An overview of studies that examined the effect of phytoplasma infections on phytochemical con-tent and other secondary metabolites of affected plants is also included. Phytoplasma diseases of medicinal and aromatic plants occur worldwide; however, the majority of reports are from Europe and southeastern Asian coun-tries. These diseases affect plant species belonging to over 70 families, mostly to Apiaceae and Asteraceae. They dif-fer considerably in geographic distribution and size of the various taxonomic groups and subgroups of the associ-ated phytoplasmas. Subgroup 16SrI-B phytoplasmas are the prevalent agents occurring mainly in Europe, North America and Asia. Phytoplasma presence induces changes in the amount and composition of secondary metabolites in diseased plants in which, however, the concentrations of valuable phytochemicals are greatly affected. An excep-tion is represented by phytoplasma diseases of periwinkle in which an accumulation of pharmaceutically impor-tant compounds occurs upon phytoplasma infections. Prospects for future research are presented and critically discussed.

Keywords: ‘Candidatus Phytoplasma’ species, 16Sr group/subgroups, phytochemicals, secondary metabolites, symptomatology, yellows diseases.

INTRODUCTION

Medicinal and aromatic plants include a broad array of wild and cultivated plants which provide raw materials for medicines, nutraceuticals, perfumes, flavors and cosmetics. These plants contain many biologically-active compounds, namely secondary metabolites, which are known as phyto-chemicals that are of great interest for their ability to pro-mote human and animal health. Currently, the demand for medicinal and aromatic plants and their derived products is greatly increasing. Among the several diseases affecting medicinal and aromatic plants, there are yellows, witches’ broom and decline diseases, associated with phytoplasma presence, which severely impair productivity, phytochemi-cal content and longevity of affected plants.

Phytoplasmas are a large group of plant-pathogenic wall-less bacteria associated with diseases in more than a thousand plant species worldwide. They are transmitted from plant to plant by phloem-feeding homopteran in-sects, mainly leafhoppers and planthoppers, less frequently psyllids, in a persistent, propagative manner (Weintraub and Beanland, 2006). Most of the phytoplasma host plants are angiosperms in which a wide range of specific and non-specific symptoms are induced (Seemüller et al., 2002; Bertaccini et al., 2014). Specific symptoms include virescence, phyllody, big bud, flower proliferation and other flower abnormalities, resulting in sterility, witches’ brooms, rosetting, internode elongation and etiolation, shortened internodes, off-season growth, and brown dis-coloration of phloem tissue. Less specific and non-specific symptoms which are most often common in woody plants, include foliar yellowing and reddening, small leaves, leaf roll, leaf curl, vein clearing, vein enlargement, vein necro-sis, premature autumn coloration, premature defoliation, poor terminal growth, sparse foliage, die-back, stunting, and decline.

Since these prokaryotes live and multiply in functional phloem sieve tube elements, they mainly impair their func-tionality. The inhibition of phloem transport along with other impaired physiological functions including reduced photosyntesis, altered secondary metabolism and plant hormone imbalance explain the symptoms exhibited by phytoplasma-infected plants (Lepka et al., 1999; Tan and Whitlow, 2001; Bertamini et al., 2002; Choi et al., 2004;

Journal of Plant Pathology (2016), 98 (3), 379-404 379 Edizioni ETS Pisa, 2016

Corresponding author: C. Marcone Fax: +39.089.969602 E-mail: [email protected]

380 Phytoplasma diseases Journal of Plant Pathology (2016), 98 (3), 379-404

Endeshaw et al., 2012; Ding et al., 2013; Vitali et al., 2013). However, the mechanisms by which phytoplasmas induce disease in plants and the different reactions of the host plants are still poorly understood. Several plant host genes, which are differentially expressed upon phytoplasma in-fections, were identified. These include genes involved in phytohormone activity, photosynthesis, carbohydrate and lipid metabolism, amino acid transport, and secondary metabolite biosynthesis genes (Pracros et al., 2006; Chen and Lin, 2011; Landi and Romanazzi, 2011; Albertazzi et al., 2009; Hren et al., 2009a, 2009b; Ding et al., 2013). In addition, a considerable number of genes that are likely to play major roles in phytoplasma-host interactions are known from the complete genome sequences of the phyto-plasmas that have been determined to date. Among these, there are genes encoding surface membrane proteins and effector (virulence) proteins (Hoshi et al., 2009; MacLean et al., 2011; Sugio et al., 2011a, 2011b; Kube et al., 2012; Neriya et al., 2014).

Taxonomically, phytoplasmas are placed in the class Mollicutes and are currently classified within the provi-sional genus ‘Candidatus Phytoplasma’ based primarily on 16S rDNA sequence analysis (IRPCM, 2004; Martini et al., 2014). Approximately 33 major phylogenetic groups were identified within the phytoplasma clade (Bertaccini et al., 2014). This figure is broadly in accordance with the number of phytoplasma groups established by restric-tion fragment length polymorphism (RFLP) analysis of PCR-amplified rDNA (Lee et al., 1998, 2000; Seemüller et al., 1998, 2002). Within the majority of the phytoplasma groups, several distinct subgroups have been delineated, based on the RFLP analysis of 16S rDNA sequences (Lee et al., 1998, 2007). To date, 40 ‘Candidatus Phytoplasma’ species have been formally described (IRPCM, 2004; Bertaccini et al., 2014; Davis et al., 2016; Fernández et al., 2016; Šafářová et al., 2016).

This review provides a literature overview of phyto-plasma diseases affecting medicinal and aromatic plants examining also the effect of phytoplasma presence on the phytochemical content of affected plants.

PHYTOPLASMA DISEASES OF MEDICINAL AND AROMATIC PLANTS

Phytoplasma diseases of medicinal and aromatic plants occur worldwide; however, the majority of the reports are from Europe and southeastern Asian countries (Table 1). Many of these diseases either were previously of unknown etiologies or mistakenly presumed to be induced by virus. Phytoplasma diseases of medicinal and aromatic plants affect plant species belonging to over 70 families, most-ly to Apiaceae and Asteraceae. They differ considerably in geographic distribution and size of the various taxo-nomic groups and subgroups of the associated phytoplas-mas. It appears that subgroup 16SrI-B phytoplasmas are

the prevalent agents occurring mainly in Europe, North America and Asia. Phytoplasmas of 16SrXII-A subgroup with a few exceptions, e.g., Artemisia scoparia witches’ broom and Salvia miltiorrhiza red leaf agents, both oc-curring in China, as well as phytoplasmas of 16SrX and 16SrXX groups are restricted to Europe, whereas phyto-plasmas of the 16SrII group are prevalent in Asia and Aus-tralia. Also, phytoplasmas of 16SrIV, 16SrXIII, 16SrXV, 16SrXXIX, 16SrXXX and 16SrXXXII groups have only been identified in diseased medicinal and aromatic plants in Asia and South America. Like other phytoplasma dis-eases, a number of diseases of medicinal and aromatic plants are associated each with genetically different phy-toplasmas, up to seven in some instances, which induce similar symptoms in a given plant host and occur either within the same geographic areas or different continents. Among these, there are dill yellows, celery yellows, peri-winkle yellows and phyllody, cirsium yellows, purple cone-flower yellows, dandelion yellows, alfalfa witches’ broom, mallow yellows, Chinaberry yellows, myrtle yellows, rose yellows, and grapevine yellows. The remaining diseases of medicinal and aromatic plants are attributed to one or a few phytoplasmas (Table 1). Also, a given plant host may be singly or doubly or multiply infected with distinctly dif-ferent phytoplasmas. The latter phenomenon is common in perennial plants, e.g., Spanish broom, eucalypt, myrtle, olive, pomegranate, rose, rosemary, lavender, whose long life spans provide opportunities to be infected by insect vectors carrying various phytoplasmas (Table 1). In many instances, wild medicinal and aromatic plants listed in Table 1 act as alternative hosts for phytoplasmas associ-ated with economically important diseases of vegetable, ornamental, and fruit crops. These include field bindweed, hedge bindweed, nettle, and lavender that are alternative hosts of the “stolbur” phytoplasma, pot marigold, dandeli-on, broad leaf plantain, wild carrot, daisy, creeping thistle, prickly lettuce and wild radish, which are alternative hosts of aster yellows phytoplasmas, howthorn and blackthorn, alternative hosts of phytoplasmas of the 16SrX group in which phytoplasmas infecting fruit trees are also enclosed.

EFFECT OF PHYTOPLASMAS ON PHYTOCHEMICAL CONTENT AND OTHER SECONDARY METABOLITES IN AFFECTED PLANTS

There are several studies over the last decade in which the effects of phytoplasma presence on phytochemical con-tent and other secondary metabolites were investigated in symptomatic plants using mainly chromatographic, spec-trometric and molecular based-methods.

St. John’s wort. The paper by Bruni et al. (2005) showed that qualitative and quantitative phytochemical changes occurred in Hypericum perforatum (St. John’s wort) ‘Zorzi’ plants infected with a 16SrVII phytoplasma (‘Candidatus

Journal of Plant Pathology (2016), 98 (3), 379-404 Marcone et al. 381

Phytoplasma fraxini’) and showing yellows and witches’ broom symptoms, in comparisons to healthy plants. The total flavonoid and naphthodianthrone contents of dis-eased plants were lower than those of healthy ones (12.55 versus 6.74 mg/g and 1.41 versus 2.29 mg/g, respectively). Among flavonoid and naphthodianthrone components, affected plants showed markedly decreased levels of ru-tin (1.96 versus 4.96 mg/g), hyperoside (2.38 versus 3.04 mg/g), isoquercitrin (1.47 versus 3.50 mg/g), amentoflavone (0.12 versus 0.39 mg/g) and pseudohypericin (1.41 versus 2.29 mg/g), whereas the levels of hypericin, quercitrin, and quercetin were not significantly reduced. In contrast, the amount of chlorogenic acid, a phenylpropane component, was higher in diseased plants that in healthy ones (1.56 ver-sus 0.77 mg/g). Moreover the yield of volatile fraction, i.e., essential oils, was drastically reduced in diseased plants (0.11 versus 0.75% in healthy plants) and these oils showed an increase in sesquiterpenes (β-caryophyllene, β-copaene, γ-amorphene, δ-elemene, α-humulene, and germacrene D), from 63.52 to 75.01%, and a significant decrease of mono-terpene hydrocarbons (from 8.63 to 4.14%) and aliphatic compounds (from 6.31 to 2.58%). Flavonoids, naphthodi-anthrones and essential oils are major phytochemicals of the H. perforatum drugs, which provide medical and/or human health benefits.

Gum weed. Grindelia robusta (gum weed) plants in-fected by the aster yellows agent ‘Ca. P. asteris’, subgroup 16SrI-B, contained less essential oils than healthy plants (0.11 versus 0.14%) (Bertaccini et al., 2008, 2011; Bellardi et al., 2009). Forty-two compounds were identified in the essential oils of both healthy and diseased plants. The levels of most of these compounds were lower in essen-tial oils of diseased plants than in those of healthy plants. However, levels of monoterpenes such as limonene (9.34 versus 4.21%), borneol (21.34 versus 15.02) e bornyl acetate (8.91 versus 4.44%) were higher in infected plants than in healthy ones.

Wild bergamot. Essential oils extracted from Mo-narda fistulosa (wild bergamot) plants showing symp-toms of virescence, yellowing and stunting, which were doubly infected with subgroup 16SrI-B and 16SrXII-A phytoplasmas, presented an increase in the amount of some monoterpenes such as α-thujene, α- and β-pinene, and β-phellandrene as well as in the amount of α-caryophyllene and a marked decrease in the amount of thymol, compared to healthy plants. However, essential oils extracted from symptomless M. fistulosa plants, which were infected by 16SrXII-A phytoplasma alone, showed only a significant increase in the thymol content and a marked decrease in the amount of monoterpenes, com-pared to healthy plants (Contaldo et al., 2011). Thymol is a major compound of plants having antibacterial and antimycotic properties such as M. fistulosa. Therefore, further sudies are needed to verify the maintainance of

the therapeutic effects of essential oils extracted from M. fistulosa plants doubly infected with the above mentioned phytoplasmas.

Hyssop. Essential oils from Hyssopus officinalis (hys-sop) plants affected by the hyssop yellows disease, which is mostly associated with a 16SrXII-A phytoplasma, showed lower contents of isopinocanfone and trans-pinocanfone than those of healthy plants. These compounds occurred at a rate of 29 and 3.92%, respectively, in diseased plants, whereas amounted to 44.7 and 6.2% respectively, in healthy plants. Conversely, the limonene content of es-sential oils from diseased H. officinalis plants was 8.87% whereas that from healthy plants was 1.86% (Bellardi and Bertaccini, 2005; Bertaccini et al., 2005).

Purple coneflower. In Echinacea purpurea (purple cone-flower) plants infected with a subgroup 16SrIX-C phyto-plasma and showing symptoms of phyllody and foliar red-dening the contents of cichoric acid and main caffeic acid derivatives were lower than those of healthy plants (15.38 versus 18.35 mg/g and 18.15 versus 20.82 mg/g, respec-tively). No significant differences between diseased and healthy plants were found in alkamide contents (Pellati et al., 2011). Diseased purple coneflower plants showed a yield of essential oils, which differed slightly from that of healthy plants (0.026 versus 0.029% in healthy plants). Sig-nificant differences between diseased and healthy plants in the components of essential oils were observed for limonene (4.4 versus 2.2% in healthy plants), cis-verbenol (5.7 versus 1.8% in healthy plants), verbenone (11.6 ver-sus 2.7% in healthy plants), carvone (2.5 versus 0.8% in healthy plants), germacrene D (8.5 versus 10.8% in healthy plants), caryophyllene oxide (3.3 versus 4.5% in healthy plants), and spathulenol (3.2 versus 4.4% in healthy plants) (Pellati et al., 2011).

Pellitory. Flavonoid contents of Parietaria officinalis and P. judaica (pellitory) plants were severely affected by 16SrXII-A phytoplasmas. Kaempeferol and isorhamne-tin compounds could not be detected in diseased plants, while they occurred at concentrations of 47.56 and 22.28 mg/100 g of dry weight, respectively, in healthy plants. Al-so, quercetin content was lower in diseased plants than in healthy ones (10.26 versus 59.43 mg/100g of of dry weight) (Bellardi et al., 2008).

Pyrethrum. Tanacetum cinerariifoliumm (pyrethrum) is a valuable source of the secondary metabolite pyrethrin, widely used for environmentally friendly insecticides. In vitro-grown plants of pyrethrum, experimentally infected with a subgroup 16SrIII-B phytoplasma, showed a marked decrease of pyrethrin I and pyrethrin II contents, com-pared to healthy in vitro-grown plants. The decrease was less pronounced in plants grown on Murashige and Skoog (MS) medium than in those on MS medium supplemented


with 3 µM IAA and 2 µM BAP (Ambrožič-Dolinšek et al., 2008).

Spanish broom. The paper by Mancini et al. (2010a) has shown that the yield of essential oils extracted from flow-ers of Spartium junceum (Spanish broom) plants affected by the spartium witches’ broom (SpaWB) disease was a little bit lower than that of healthy plants (0.004 versus 0.015%). Also, essential oils from healthy and diseased plants were qualitatively different. Substantial amounts of sesquiterpenes and a marked decrease in the amount of n-alkanes and aliphatic compounds occurred in the es-sential oils from flowers of diseased plants. Sesquiterpenes were detected only in the essential oils of diseased plants. Approximately one third of the essential oils of infected plants was represented by six sesquiterpenes (30.5%). Among them, five (25.5%) were oxygenated compounds with τ-cadinol (19.2%) as the main component. In both healthy and diseased plants, the main constituents were n-alkanes, which occurred at rate of 55.2 and 38.8%, respectively: tricosane (14.4%), pentacosane (7.3%) and heptacosane (4.6%) were the main components in healthy plants, whereas tricosane (5.5%), nonacosane (5.3%) and triacontane (5.0%) were prevalent in diseased plants. Ali-phatic compounds occurred at a rate of 4.5% in diseased plants, whereas they amounted to 18.7% in healthy plants (Mancini et al., 2010a). Great differences also occurred in the content of alkaloid compounds between healthy and diseased Spanish broom plants: its content was con-siderably higher in samples of infected plants. Seven dif-ferent alkaloids were identified only in diseased plants. These compounds included N-methylcytisine, its isomer, N-formylcytisine and a hydroxy-substituted derivative of sparteine. Four alkaloids including hydroxy-derivatives of cytisine and anagyrine were shared by both healthy and diseased plants. All identified alkaloids were of the qui-nolizidine type (Mancini et al., 2010b).

Periwinkle. Catharanthus roseus (periwinkle) is a me-dicinal plant of pharmaceutical interest for its capacity to biosynthesize more than 130 alkaloids, most of which have pharmacological activities. Among these, there are terpenoid indole alkaloids, including vinblastine and vin-cristine, which are used for their antineoplastic activity in the treatment of many cancers, and ajmalicine and ser-pentine, which are used as antihypertensive and sedative, respectively (Zhou et al., 2009; Pan et al., 2016). However, terpenoid indole alkaloids are produced in periwinkle only in very small amounts and usually have some chiral cen-tres. They are therefore difficult to synthesize chemically, making expensive their large scale production (Zhou et al., 2009). Phytoplasma infections in periwinkle cause an increase of metabolites related to the biosynthetic path-ways of terpenoid indole alkaloids (loganic acid, secolo-ganic, vindoline) and phenylpropanoids (chlorogenic acid and polyphenols) (Choi et al., 2004; Favali et al., 2004).

In particular, it has been shown that as a consequence of the infection of periwinkle plants with clover phyllody phytoplasma, (16SrIII-B subgroup), the content of terpe-noid indole alkaloids such as vindoline, ajmalicine, ser-pentine, vinblastine and vincristine increased. The total alkaloid content of dry weight was 550.96 μg/g, whereas that of healthy plants was 452.48 μg/g (Favali et al., 2004). However, the level reached by each compound in diseased plants greatly differed among the plant organs. Vinblastine reached its highest concentration in the roots of infect-ed plants (148.44 μg/g), whereas it was not detectable or present at very low level (< 0.05 μg/g) in those of healthy plants. The content of vindoline (precursor of vinblas-tine) increased in diseased leaves (222.00 versus 131.90 μg/g of healthy plants) and stems (13.22 versus < 0.05 μg/g of healthy plants) but decreased in diseased roots (< 0.05 versus 142.95 μg/g of healthy plants). Also, phytoplasma infections led to a significant, two- to four-fold increase in chlorogenic acid in periwinkle, compared to healthy plants (Choi et al., 2004). Work by Srivastava et al. (2014) has shown that in flowers of periwinkle plants infected with a 16SrI-B phytoplasma the contents of vindoline, catharanthine, and vincristine plus vinblastine were two-, ten-, and four-fold higher, respectively, than those of healthy flowers, whereas the contents of serpentine plus ajmalicine were lower than those of healthy flowers. How-ever, in the leaves there were no significant differences between healthy and infected plants. Also, several genes including some involved in mevalonate pathway of the ter-penoid indole alkaloid biosynthesis system, namely geranyl geranyl pyrophosphate synthase, geraniol 10 hydroxylase, desacetoxyvindoline-4-hydroxylase, strictosidine synthase, secologanin synthase, and deacetylvindoline-4-O-acetyl-transferase were expressed at higher levels in flowers of diseased periwinkles than in those of healthy plants (Sriv-astava et al., 2014). Over the last few years, many attempts have been made to boost the yield of terpenoid indole al-kaloids in periwinkle. Despite the progress made, they still cannot be successfully produced on an industrial scale, owing to several biological and technological constraints. The various means by which terpenoid indole alkaloid ac-cumulation can be increased include biotic elicitors such as phytoplasmas (Zhou et al., 2009). Therefore, phytoplas-ma infections in periwinkle, in spite of the detrimental effects on plant host, could be considered as beneficial from the pharmaceutical point of view.

Grapevine. Grapevine yellows (GY) are diseases of Vi-tis vinifera (grapevine) characterized by similar symptoms but differing in associated phytoplasmas and epidemiol-ogy. Infections of grapevine plants by the “stolbur” agent ‘Ca. P. solani’, subgroup 16SrXII-A, during the early stages of shoot lignification induced a significant accumulation of several phenolic compounds including hydroxycinnamic acid, monolignol derivatives, flavanone, ellagic acid de-rivatives, flavonols, flavanols and stilbenoids (Rusjan and


Mikulic-Petkovsek, 2015). The significantly higher con-centration of hydroxycinnamic acid, monolignol deriva-tives and flavanone in shoots of diseased plants, compared to healthy ones, is a clear evidence that phenylpropanoid and cinnamate/monolignol pathways were altered after the phytoplasma infections, thereby leading to lack or incom-plete shoot lignification (Rusjan and Mikulic-Petkovsek, 2015). ‘Ca. P. solani’ infections also induced an accumu-lation, mainly during the early phenological stages, in the leaves of diseased grapevine plants, of total phenols, hydroxycinnamic acid, flavonols and flavanols. Further-more, infected leaves showed a marked activity increase of phenylalanine ammonia liase (PAL), chalcone synthase (CHS), chalcone isomerase (CHI), flavanone 3-hydroxylase (FHT) and polyphenoloxidase (PPO), and a decrease of peroxidase (POD), compared to healthy plants, through-out the whole growing season. In addition, chlorophyll and carotenoid contents were considerably reduced in infected leaves (Bertamini et al., 2002; Landi and Romanazzi, 2011; Endeshaw et al., 2012; Rusjan et al., 2012a). Investigations on the accumulation of phenolic compounds in berries of “stolbur”-affected grapevine plants, at three different phenological stages such as veraison, berry softening and berries ripe for harvest, showed that the total phenolic content of berries from diseased plants was higher than that of berries from healthy plants at veraison and berry softening, whereas the opposite was true at stage of berries ripe for harvest (Rusjan et al., 2012b). Hydroxycinnamic acids and flavanols content was higher in berries from diseased plants, compared to healthy ones only at berry softening stage, whereas the content of flavonols in ber-ries from diseased plants proved to be considerably lower than that of healthy plants in each of the three pheno-logical stages (Rusjan et al., 2012b). Decreased contents of phenolic compounds especially flavonols in berries ripe for harvest greatly impair quality of grape and wine and their antioxidant properties (Rusjan et al., 2012b). Grape berries are valuable nutraceuticals mainly due to the pres-ence of different types of flavonoids such as flavonols and anthocyanins and are widely used in pharmaceutical and cosmetic industries (Rusjan et al., 2008). Several genes involved in secondary metabolism were differentially ex-pressed in grapevine plants upon ‘Ca. P. solani’ infection. Among these, PAL, CHS, FHT, dihydroflavonol 4-reduc-tase, leucoanthocyanidin dioxygenase, flavonol synthase, beta-carotene hydroxylase and carotenoid cleavage di-oxygenase 1 were significantly up-regulated in diseased plants, compared with healthy ones (Albertazzi et al., 2009; Hren et al., 2009a; Landi and Romanazzi, 2011). Genes encoding Myb transcription factors were down-regulated in highly susceptible grapevine cv Chardonnay plants by ‘Ca. P. solani’ infections. Since Myb transcription factors are essential for lignin biosynthesis, down-regulation of these genes may be another factor accounting for the lack of, or incomplete, shoot lignification in diseased grape-vine plants (Albertazzi et al., 2009; Hren et al., 2009a).

Proteomic studies also revealed that a number of proteins were differentially expressed in grapevine cv Nebbiolo plants infected by the “flavescence dorée” phytoplasma, a member of the 16SrV group, compared to healthy plants. Among the differentially expressed proteins, a chalcone isomerase was strongly up-regulated in affected plants (Margaria and Palmano, 2011; Margaria et al., 2013).

CONCLUSIONS AND OUTLOOK

This review is the first to compile a comprehensive list of phytoplasmas and phytoplasma diseases affecting me-dicinal and aromatic plants as reported in the last years. Newly discovered diseases of such kind of plants are in-creasingly being attributed to phytoplasma presence. This rapid increase may be related to the fact that the use of chemical insecticides to control insect vectors of phyto-plasmas is not allowed on medicinal and aromatic plants, whereas insect control and/or management by other, environmentally-friendly means is not fully satisfactory. Phytoplasma diseases of medicinal and aromatic plants severely reduce yield and quality of crops and longevity of plants. Changes in the composition of secondary me-tabolites occurring in diseased plants are related to the role of phytoplasmas in triggering plant defense responses in which, however, the levels of valuable phytochemicals are greatly affected. An exception is represented by peri-winkle in which an accumulation of pharmaceutically important compounds such as vinblastine and vincristine occurs upon phytoplasma infection. Also, because the beneficial properties of medicinal and aromatic plants can be related to combinations of phytochemicals act-ing collectively or synergistically, the alteration of a single phytochemical component could potentially affect the ef-ficacy of the plant material (Briskin, 2000). Therefore, any variation of the chemical composition of medicinal and aromatic plants would need to be investigated by phar-macological studies aimed at evaluating the plant mate-rial effectiveness. The impact of phytoplasma infections on medicinal and aromatic plants should be taken into account in promoting good agricultural pratices for culti-vation and propagation of these plants and for the develop-ment of effective and standardised phytopharmaceutical and nutraceutical products. Moreover, further studies are needed to acquire knowledge on several other aspects of the above mentioned diseases including disease manage-ment, phytoplasma-insect vector relationships and role of phytoplasmas in eliciting secondary metabolites which can be pharmaceutically important.


Table 1. Phytoplasma diseases of medicinal and aromatic plants.

Family Genus/species

Common name

Disease Phytoplasma(s) associateda

Geographic distribution

Reference

AcanthaceaeAndrographis paniculata Kalmegh Kalmegh witches’

broom16SrII-D India Saeed et al., 2015

AgavaceaeAgave tequilana Blue agave Agave lethal

yellowing16SrIV Mexico González-Pacheco et al., 2014

AmaranthaceaeAchyranthes aspera Prickly chaff flower Achyranthes yellows 16SrI India Raj et al., 2009aAmaranthus retroflexus Common amaranth Unnamed 16SrV-B, 16SrXII-A China, Czech Republic,

ItalyFialová et al., 2009; Yang et al., 2011; Mori et al., 2015

Amaranthus hypochondriacus

Grain amaranth Unnamed 16SrII, 16SrIII Mexico Rojas-Martinez et al., 2009; Ochoa-Sanchez et al., 2009; Pérez-López et al., 2016

Amaranthus cruentus Grain amaranth Unnamed 16SrII Mexico Ochoa-Sanchez et al., 2009Amaranthus spp. Amaranth Amaranth yellows 16SrI-B, 16SrII, 16SrXII-A USA, India, Turkey Borth et al., 2006; Arocha et al., 2009;

Özdemir et al., 2009Chenopodium album Lambsquarters Chenopodium

yellows16SrII, 16SrIII, 16SrXII-A Europe, Turkey Tolu et al., 2006; Özdemir et al., 2009;

Safarova et al., 2011; Mori et al., 2015

AnacardiaceaePistacia palaestina Terebinth Unnamed 16SrIX-C Lebanon Casati et al., 2016

Apiaceae Amni majus Queen Anne’s lace Queen Anne’s lace

yellows16SrI-B Canada Wang and Hiruki, 2005

Anethum graveolens Dill Dill yellows 16SrI-A, -B, -C, 16SrV-A, 16SrXII-A

USA, Italy Boccardo et al., 2002; Lee et al., 2003, 2004

Anthriscus cerefolium Curled chervil Anthriscus reddening N.D. Canada Hwang et al., 1998Apium graveolens Celery Celery yellows 16SrI-A, -B, 16SrII-E,

16SrIII; 16SrXII-A, -BEurope, North America, Japan, Australia, New Zealand

Seemüller et al., 1998; Tran-Nguyen et al. 2003; Lee et al., 2004; Carraro et al., 2008; Fialová et al., 2009; Wang and Hiruki, 2005; Ivanović et al., 2011; Liefting et al., 2011; Maejima et al., 2014

Apium graveolens Celery Celery stunting and phyllody

16SrI-C Czech Republic Fránová and Špak, 2013

Bupleurum falcatum Chai hu Chai hu yellows 16SrI-A Canada Chang et al., 2004Carum carvi Caraway Caraway yellows N.D. Canada Hwang et al., 1997aCarum carvi Caraway Caraway yellows 16SrXII-A Serbia Pavlović et al., 2014aCoriandrum sativum Coriander Coriander yellows 16SrI-A, -B USA Lee et al., 2004Daucus carota Carrot Carrot yellows 16SrI-A, -B, 16SrIX,

16SrXII-ANorth America, Europe, Japan

Chang et al., 2004; Lee et al., 2004; Maejima et al., 2014; Marchi et al., 2015

Eryngium alpinum Queen of the Alps Eryngium yellows 16SrI-M Lithuania Samuitiene et al., 2006, 2007Foeniculum vulgare Fennel Fennel phyllody 16SrII India Bhat et al., 2008Levisticum officinale Lovage Levisticum yellows 16SrXII-A Serbia Pavlović et al., 2014aPetroselinum spp. Parsley Parsley yellows 16SrI-A, -B, 16SrXII-A Canada, Serbia Khadhair et al., 1998; Lee et al, 2003,

2004; Wang and Hiruki, 2005; Mitrović et al., 2013

Pimpinella anisum Anise Unnamed 16SrI Canada Olivier et al., 2009Trachyspermum ammi Ajwain Ajwain yellows 16SrVI India Samad et al., 2011a

ApocynaceaeCatharanthus roseus Periwinkle Mexican periwinkle

virescence16SrXIII-A (‘Ca. P. hyspanicum)

Mexico Davis et al., 2016

Catharanthus roseus Periwinkle Malaysian periwinkle virescence

16SrXXXII-A (‘Ca. P. malaysianum’)

Asia Nejat et al., 2013

Catharanthus roseus Periwinkle Periwinkle yellows and phyllody

16SrI, 16SrII, 16SrIII, 16SrVI, 16SrIX, 16SrXII

Europe, Americas, Asia Seemüller et al., 1998; Lee et al., 2000, 2004, 2012; Torres et al., 2004; Ayman et al., 2010; Chen et al., 2011; Galdeano et al., 2013; Mitrović et al., 2013; Caicedo et al., 2015; Khanna et al., 2015; Pérez-López et al., 2016

Periploca aphylla Milk broom Periploca aphylla witches’ broom

16SrXXX (‘Ca. P. tamaricis’)-related

Iran Faghihi et al., 2010

Calotropis gigantea Crown flower Calotropis yellows 16SrVI India Priya et al., 2010Tylophora asthmatica Indian ipecac Tylophora asthmatica

little leaf16SrII-D India Madhupriya et al., 2014

ArecaceaeRoystonea regia Royal palm Royal palm yellow

decline16SrI Malaysia Naderali et al., 2015

AsclepiadaceaeAsclepias fruticosa (syn. Gomphocarpus fruticosa)

Swan plant Swan plant yellows 16SrXII-B New Zealand Liefting et al., 2011


Asclepias physocarpa (syn. Gomphocarpus physocarpus)

Balloonplant Balloonplant yellows 16SrI-B, 16SrXII-A Italy, Iran d’Aquilio et al., 2002; Bellardi and Bertaccini, 2005; Bertaccini et al., 2006; Babaie et al., 2007

Asclepias syriaca Milkweed Milkweed yellows 16SrIII-F North America Lee et al., 2000

AsparagaceaeAsparagus officinalis Asparagus Asparagus stem

fasciation16SrI-B Czech Republic Fránová and Petrzik, 2010

Asparagus officinalis Asparagus Asparagus yellows 16SrI-A Canada Wang and Hiruki, 2005

Asteraceae Arnica montana Mountain arnica Arnica virescence 16SrIII-B Serbia Pavlovic et al., 2012aArtemisia scoparia Redstem

wormwoodArtemisia scoparia witches’ broom

16SrXII-A China Yu et al., 2016

Artemisia vulgaris Mugwort Artemisia yellows 16SrXII-A Italy Mori et al., 2015Bellis perennis Daisy Daisy yellows 16SrI-A, 16SrIII-J Lithuania, Argentina Samuitiene et al., 2007; Galdeano et al.,

2013Calendula officinalis Pot marigold Calendula virescence

and phyllody16SrI-B, 16SrII, 16SrXII-A Italy, Serbia, Canada,

IranHwang et al., 1977b; Marcone et al., 2000; Lee et al., 2004; Wang and Hiruki, 2005; Esmailzadeh-Hosseini et al., 2011; Pavlovic et al., 2014b

Calendula arvensis Field marigold Unnamed 16SrII-E Italy Tolu et al., 2006Centaurea solstitialis Yellow starthistle Centaurea solstitialis

virescence16SrVI Italy Faggioli et al., 2004

Cichorium intybus Common chicory Chicory phyllody 16SrII-E, 16SrIX, 16SrXII-A

Italy, Serbia, Australia Tran-Nguyen et al. 2003; Marcone, 2011; Ermacora et al., 2013; Pavlovic et al., 2014c

Cirsium arvense Creeping thistle Cirsium yellows and stunting

16SrXI-E (‘Ca. P. cirsii’) Czech Republic Šafářová et al., 2016

Cirsium arvense Creeping thistle Cirsium yellows 16SrI-M., 16SrIII, 16SrIX, 16SrXII-A

Europe Seemüller et al., 1998; Jomantiene et al., 2000, 2002; Lee et al., 2004; Bellardi and Bertaccini, 2005; Fialová et al., 2009; Marchi et al., 2015

Cirsium arvense Creeping thistle Cirsium multiple inflorescence

16SrIII-B Serbia Rančić et al., 2005; Jakovljević et al., 2015

Cynara scolymus Artichoke Artichoke degeneration

16SrXII Spain Battle et al., 2007

Dittrichia viscosa (syn. Inula viscosa)

False yellowhead Unnamed 16SrIX-C Lebanon Casati et al., 2016

Echinacea angustifolia Narrow-leaved purple coneflower

Purple coneflower yellows

16SrXII-A Serbia Pavlović et al., 2014a

Echinacea purpurea Purple coneflower Purple coneflower yellows

16SrI-A, -B, -C; 16SrIII-B; 16SrIX-C; 16SrXII-A

North America, Europe Khadhair et al., 1997; Hwang et al., 1977c; Lee et al., 2004, 2008; Wang and Hiruki, 2005; Bertaccini et al., 2009; Radisek et al., 2009; Fránová et al., 2009, 2013; Pavlović et al., 2011; Pellati et al., 2011

Echinacea purpurea Purple coneflower Purple coneflower witches’ broom

16SrII-A Taiwan Tseng et al., 2012

Echinacea pallida Pale purple coneflower

Pale purple coneflower witches’ broom

16SrII-D Australia Pearce et al., 2011

Echinops spinosissimus Hink Echinops witches’ broom

16SrIX Oman Al-Subhi et al., 2007

Grindelia robusta Gum weed Unnamed 16SrI-B Italy Bertaccini et al., 2008, 2011; Bellardi et al., 2009

Guizotia abyssinica Niger seed Niger seed phyllody 16SrI, 16SrII Iran, Australia Seemüller et al., 1998; Vaali et al., 2011Ixeridium chinense China ixeris China ixeris

fasciation16SrI-C China Li et al., 2013

Lactuca serriola Prickly lettuce Prickly lettuce yellows

16SrI-B, 16SrIX-C Iran, Lebanon Babaie et al., 2007; Salehi et al., 2007; Sichani et al., 2014; Casati et al., 2016

Leucanthemum vulgare (syn. Chrysanthemum vulgare)

Oxeye daisy Oxeye daisy phyllody 16SrIII-B Italy Firrao et al., 1996

Matricaria chamomilla (syn. Chamomilla recutita)

Chamomile Chamomile witches’ broom

16SrXII-A Serbia Pavlović et al., 2014a

Parthenium hysterophorus Whitetop weed Parthenium witches’ broom

16SrI, 16SrII Asia Raj et al., 2008a; Li et al., 2011a; Ahmad et al., 2015

Sonchus asper Grespino thorny Grespino thorny yellows

16SrI-A USA Lee et al., 2004

Sonchus spp. Sowthistle Unnamed 16SrI, 16SrIX, 16SrXII-A Europe, North America Borth et al., 2006; Khadhair et al., 2008; Marchi et al., 2015; Mori et al., 2015

Stevia rebaudiana Honey leaf plant Stevia decline 16SrXXIV-A India Samad et al., 2011bTagetes erecta Marigold Marigold phyllody 16SrI-B, 16SrXII-A Americas, Europe, Asia,

TurkeyLoi et al., 1998; Rojas-Martínez et al., 2003; Raj et al., 2011a; Alp et al., 2016

Tagetes patula French marigold Marigold yellows 16SrI-A, -B Americas, Europe, Asia Bellardi and Bertaccini, 2005; Wang and Hiruki, 2005; Babaie et al., 2007; Sichani et al., 2014

Table 1 (continued). Phytoplasma diseases of medicinal and aromatic plants.


Tanacetum cinerariifoliumm

Pyrethrum Pyrethrum yellows 16SrI-A Canada Wang and Hiruki, 2005

Tanacetum parthenium Feverfew Feverfew yellows 16SrI-B Canada, Iran Wang and Hiruki, 2005; Babaie et al., 2007

Taraxacum officinale Dandelion Dandelion yellows 16SrI, 16SrII-E, 16SrIII, 16SrIX, 16SrXII-A

Europe, Americas Seemüller et al., 1998; Jomantiene et al., 2002; Lee et al., 2004; Wang and Hiruki, 2005; Hodgetts et al., 2009; Marcone, 2011; Mori et al., 2015

Tussilago farfara Coltsfoot Tussilago reddening 16SrXII-A Italy Romanazzi et al., 2009; Marchi et al., 2015

BalsaminaceaeImpatiens balsamina Rose balsam Rose balsam phyllody

and virescence16SrI-D China Chang et al., 2011; Li et al., 2011b

BerberidaceaeBerberis thunbergii atropurpurea

Red barberry Berberis phyllody 16SrV-B China Li et al., 2010a

BetulaceaeAlnus glutinosa Black alder Alder yellows 16SrV-C Europe Marcone, 2015

BignoniaceaeTabebuia heterophylla Tabebuia Tabebuia witches’

broom16SrIX Puerto Rico Caicedo et al., 2015

BrassicaceaeBrassica spp. Brassicas Brassica phyllody and

virescence16SrI, 16SrIII, 16SrIX, 16SrXII-A

Europe, Americas, Asia Marcone et al., 2000; Lee et al., 2004, 2012; Salehi et al., 2011; Trkulja et al., 2011; Mou et al., 2012; Rappussi et al., 2012; Eckstein et al., 2013

Eruca sativa Salad rocket Eruca sativa phyllody 16SrI-B Iran Esmailzadeh Hosseini et al., 2015aRaphanus raphanistrum Wild radish Wild radish yellows 16SrI-B Italy Marcone et al., 2000; Lee et al., 2004

CannabaceaeCannabis sativa Hemp Hemp witches’

broom16SrI, 16SrXIV-A India, Iran Raj et al., 2008b; Sichani et al., 2014;

Chaube et al., 2015a

ConvolvulaceaeCalystegia sepium Hedge bindweed Bindweed yellows 16SrXII-A Europe Langer and Maixner, 2004; Fialová et al.,

2009;Coccinia grandis Ivy gourd Ivy gourd yellows 16SrXIV Thailand Sunpapao, 2014Convolvulus arvensis Field bindweed Bindweed yellows 16SrX, 16SrI, 16SrXII-A,

16SrII, 16SrIXEurope, Israel Marcone et al., 1997a; Seemüller et al.,

1998; Langer and Maixner, 2004; Bellardi and Bertaccini, 2005; Fialová et al., 2009; Murolo et al., 2010; Dafny Yelin et al., 2015; Marchi et al., 2015; Mori et al., 2015

Convolvulus arvensis Field bindweed Bindweed yellows 16SrXII-H (‘Ca. P. convolvuli’)

Italy, Serbia, Bosnia and Herzegovina, Germany

Martini et al., 2012

CaprifoliaceaeSambucus nigra Black elder Sambucus decline 16SrXII-A, -E Italy, Germany Lederer and Seemüller, 1991; Filippin et

al., 2008Sambucus racemosa European red elder Sambucus decline N.D. Germany Lederer and Seemüller, 1991

CaryophyllaceaeSaponaria officinalis Common soapwort Saponaria yellows 16SrI-M, 16SrXII-A Serbia, Lithuania Lee et al., 2004; Josic et al., 2013Silene orientalis Campion Silene virescence 16SrI-B, 16SrXII-A Lithuania Samuitiene et al., 2006, 2007; Berger et

al., 2009

CornaceaeCornus sanguinea Common dogwood Common dogwood

yellows16SrXII-A, -E Italy, France Jarausch et al., 2001; Filippin et al., 2008

CrassulaceaeRhodiola rosea Golden root Rhodiola yellows 16SrI-B Canada Hwang et al., 2009

CucurbitaceaeMomordica charantia Bitter gourd Bitter gourd little leaf

and phyllody16SrI-B, 16SrII Myanmar, Indonesia,

ThailandSdoodee et al., 1999; Davis et al., 2003; Win et al., 2014

Lagenaria siceraria Bottle gourd Lagenaria yellows 16SrIII-Y Brazil Montano et al., 2015aSicana odorifera Sikana Unnamed 16SrIII Brazil Montano et al., 2007

CupressaceaeJuniperus occidentalis Western juniper Juniper witches’-

broom16SrIX-F USA (Oregon) Davis et al., 2010

ElaeagnaceaeElaeagnus angustifolia Russian olive Russian olive witches’

broom16SrI Iran Rashidi et al., 2010

EricaceaeVaccinium corymbosum Highbush

blueberryBlueberry stunt 16SrI-E, 16SrXII-A Canada, Serbia Pavlović et al., 2014a; Arocha Rosete et

al., 2015



Vaccinium macrocarpon American cranberry

Cranberry false-blossom

16SrIII-Y USA Lee et al., 2014

Vaccinium myrtillus Blueberry Blueberry stunt and witches’ broom

16SrI-E, 16SrIII, 16SrIX-E USA, Europe Seemüller et al., 1998; Valiunas et al., 2004; Lee et al., 2012; Bagadia et al., 2013

EuphorbiaceaeAcalypha indica Indian nettle Acalypha witches’

broomN.D. India Singh and Gupta, 2011; Singh et al.,

2013a; Singh and Upadhyaya, 2015Mallotus japonicus Food wrapper plant Mallotus witches’

broom16SrI-B Korea Han et al., 2014

Mercurialis annua Annual mercury Annual mercury yellows

16SrIII-B, 16SrXII-A Italy Bellardi and Bertaccini, 2005; Marchi et al., 2015

Sapium sebiferum Chinese tallow tree Chinese tallow tree yellows

16SrIII-Y China Gao et al., 2015a

FabaceaeCassia italica Italian senna Cassia witches’

broom16SrXXIX-A (‘Ca. P. omanense’), 16SrIX

Oman Khan et al., 2007a; Al-Saady et al., 2008

Crotalaria pallida Smooth rattlepod Crotalaria little leaf 16SrII-D India Yadav et al., 2016Dimorphandra spp. “Fava d’anta” tree Dimorphandra

witches’ broom16SrXIII Brazil Montano et al., 2015b

Galega officinalis Galega Galega yellows 16SrIII-J, 16SrVII-A, 16SrX

Chile, Italy Bellardi and Bertaccini, 2005; Longone et al., 2011; Zamorano et al., 2015

Medicago sativa Alfalfa Alfalfa witches’ broom

16SrI, 16SrII, 16SrIII-B, 16SrV-B, 16SrVI, 16SrVII-C, 16SrXII-A

Europe, Americas, Asia, Australia

Seemüller et al., 1998; Peters et al., 1999; Khan et al., 2002; Pilkington et al., 2003; Lee et al., 2004; Conci et al., 2005; Wang and Hiruki, 2005; Hodgetts et al., 2009; Li et al., 2012; Starović et al., 2012; Esmailzadeh Hosseini et al., 2015b

Mimosa pudica Sensitive plant Mimosa yellows 16SrI Indonesia Boa et al., 2010Senna surattensis Sunshine tree Sunshine tree stem

fasciation16SrV, 16SrXII China Wu et al., 2012

Sophora alopecuroides Sophora root Sophora root yellows 16SrI, 16SrVI-D China, Iran Chen et al., 2013; Zibadoost and Rastgou, 2016

Sophora japonica Chinese scholar tree

Sophora japonica witches’ broom

16SrV-B China Duduk et al., 2010; Yu et al., 2012; Ren et al., 2014

Spartium junceum Spanish broom Spartium witches’ broom

16SrX-D (‘Ca. P. spartii’), 16SrV-C

Italy, Spain Marcone et al., 1996a, 2004a; Torres et al., 2002; Spallino et al., 2013; Contaldo et al., 2015

Tephrosia purpurea Wild indigo Tephrosia purpurea witches’ broom

16SrII-M India Yadav et al., 2014

Trigonella foenum greacum Fenugreek Fenugreek yellows 16SrXII-A Serbia Pavlović et al., 2014a

GentianaceaeCentaurium erythraea Centaury Centaury virescence 16SrI-B Italy Paltrinieri et al., 2016Enicostemma axillare Indian whitehead Enicostemma

witches’ broom16SrII-C India Abirami et al., 2012

GeraniaceaeGeranium purpureum Little-robin Unnamed 16SrIX-C Lebanon Casati et al., 2016

GrossulariaceaeRibes rubrum Red currant Unnamed 16SrI-C Czech Republic Přibylová et al., 2011

HypericaceaeHypericum perforatum St. John’s wort St. John’s wort

decline16SrVII, 16SrXII-A Italy, Serbia Paltrinieri et al., 2002; Bellardi and

Bertaccini, 2005; Bruni et al., 2005; Pavlović et al., 2012b

Hypericum barbatum St. John’s wort St. John’s wort decline

16SrXII-A Serbia Pavlović et al., 2012b

LamiaceaeHyssopus officinalis Hyssop Hyssop yellows 16SrXII-A, 16SrI Italy Bellardi and Bertaccini, 2005; Bertaccini

et al., 2005Lavandula spp. Lavender Lavandula yellows 16SrXII-A, 16SrI, 16SrVI Europe, Israel Seemüller et al., 1998; Marzachì et

al., 1999; Langer and Maixner, 2004; Weintraub et al., 2007; Gaudin et al., 2011

Melissa officinalis Lemon balm Melissa yellows 16SrXII-A Serbia Pavlović et al., 2014aMentha×piperita Peppermint Peppermint stunt 16SrXII-A Serbia Mitrović et al., 2016Monarda fistulosa Wild bergamot Monarda yellows 16SrI-B, 16SrII, 16SrXII-A Canada, Italy Khadhair et al., 1997; Hwang et al., 1977d;

Wang et al., 1998; Wang and Hiruki, 2005; Bellardi et al., 2011; Contaldo et al., 2011

Ocimum basilicum Basil Basil little leaf 16SrI Cuba Arocha et al., 2006Origanum vulgare Oregano Oregano reddening 16SrXII-A Serbia Pavlović et al., 2014aRosmarinus officinalis Rosemary Rosemary yellows 16SrXII-A, 16SrI-B Italy Contaldo et al., 2012Salvia miltiorrhiza Red sage Salvia miltiorrhiza

red leaf16SrXII-A China Yang et al., 2016a

Salvia sclarea Clary sage Clary sage yellows 16SrX Italy Bellardi and Bertaccini, 2005



Thymus vulgaris Thyme Unnamed 16SrXII-A Spain Battle et al., 2000

LauraceaeCinnamomum cassia Chinese cinnamon Cinnamon yellow leaf 16SrII China Yang et al., 2016b

LiliaceaeAllium altyncolicum Unknown Unnamed 16SrI-B Czech Republic Navrátil et al., 2000Allium ampeloprasum Leek Leek proliferation 16SrI-B, 16SrXII-A Czech Republic, Italy Bertaccini et al., 1999Allium cepa Onion Onion yellows 16SrI-B, -A, -L, -M; 16SrVI Italy, Japan, USA,

Canada, Lithuania, Pakistan

Marcone et al., 2000; Khadhair et al., 2002; Lee et al., 2003, 2004; Jomantiene et al., 2010; Maejima et al., 2014; Ahmad et al., 2015

Allium fistulosum Welsh onion Onion yellows 16SrI-B Japan Maejima et al., 2014Allium sativum Garlic Garlic decline 16SrIII-J Argentina, Canada Conci et al., 1998; Khadhair et al., 2002;

Galdeano et al., 2013

MalvaceaeAbelmoschus esculentus Okra Okra bunchy top 16SrI, 16SrV, 16SrXII Mauritius, India,

PakistanFahmeed et al., 2009; Gungoosingh-Bunwaree et al., 2011; Kumar et al., 2012a

Hibiscus rosa-sinensis Chinese hibiscus Hibiscus yellows and little leaf

16SrI India Chaturvedi et al., 2010

Hibiscus rosa-sinensis Chinese hibiscus Hibiscus witches’ broom

16SrXV-A (‘Ca. P. brasiliense’), 16SrXII

Brazil Montano et al., 2001, 2011a

Hibiscus sabdariffa Roselle Roselle phyllody 16SrI, 16SrV-D India, Taiwan Biswas et al., 2013a; Tseng et al., 2014Malva sylvestris Mallow Mallow yellows 16SrI-B, 16SrV-E,

16SrIX-C, 16SrXII-AItaly, France, Lebanon Seemüller et al., 1998; Jarausch et al.,

2001; Mori et al., 2015; Casati et al., 2016Malvastrum coromandelianum

False mallow Unnamed 16SrII South America Pérez-López et al., 2016

Sida cordifolia Flannel weed Sida little leaf 16SrII-C Australia, Mali Davis et al., 1997a; Marzachì et al., 2009

MeliaceaeMelia azedarach Chinaberry Chinaberry yellows 16SrI-B, 16SrIII-J,

16SrXIII-CBolivia, Vietnam Harrison et al., 2003, 2006

Melia azedarach Chinaberry Chinaberry yellows 16SrIII-B, 16SrXIII-C (‘Ca. P. meliae’)

South America Galdeano et al., 2013; Pérez-López et al., 2016; Fernández et al., 2016

Melia azedarach Chinaberry Melia witches’ broom 16SrI-B China, Korea Han et al., 2015Toona ciliata Toon Toon yellows 16SrI-B India Ayman et al., 2010

MenispermaceaeTinospora cordifolia Heart-leaved

moonseedTinospora flat stem N.D. India Somashekhara Achar et al., 2015

Corchorus capsularis White jute White jute little leaf 16SrV India Biswas et al., 2013bCorchorus olitorius Jute Jute phyllody 16SrII-D Turkey Cagirgan et al., 2014; Özdemir and

Cagirgan, 2015

MoraceaeBrussonetia papyrifera Paper mulberry Paper mulberry

yellows16SrΙ-B China Mei et al., 2016

Streblus asper Toothbrush tree Streblus asper little leaf

16SrI India Maurya et al., 2014

MyrtaceaeEucalyptus spp. Eucalypt Eucalyptus little leaf 16SrV, 16SrI-B, -C, 16SrII Italy, India, China,

Sudan, Iran, BrazilMarcone et al., 1996b, 1997b; Camele et al., 1999; de Souza et al., 2015; Marcone, 2015; Salehi et al., 2016a

Ugni molinae Murta Murta witches’ broom

16SrIII-J, 16SrV-A, 16SrVII-A

Chile Arismendi et al., 2010, 2011a, 2011b

Myrtus communis Myrtle Myrtle yellows 16SrI-B, 16SrII-F, 16SrIII-B, 16SrV-A, 16SrX-A, 16SrXII-A

Italy Camele et al., 1999; Garau et al., 2005; Prota et al., 2007

NyctaginaceaeMirabilis jalapa Four o’clock flower Mirabilis little leaf 16SrII India, Israel Sobolev et al., 2007; Weintraub et al.,

2007; Kumar et al., 2012b

OleaceaeJasminum sambac Arabian jasmine Arabian jasmine

witches’ broom16SrI, 16SrII, 16SrXI-B Italy, Oman, India Marzachì et al., 1999; Al-Zadjali et al.,

2007; Madhupriya et al., 2015aLigustrum ovalifolium California privet Ligustrum witches’

broom16SrII Turkey Çağlar and Elbeaino, 2013

Olea europaea Olive Olive witches’ broom 16SrI-B, -C, 16SrIII, 16SrV, 16SrXII-A

Italy, Spain, Iran Seemüller et al., 1998; Font et al., 1998; Marzachì et al., 1999; Pasquini et al., 2000; Lee et al., 2004; Ahangaran et al., 2006; Çağlayan et al., 2011

OnagraceaeOenothera biennis Evening primrose Oenothera reddening

and virescence16SrXII-A, 16SrI-B Serbia, USA Hwang et al., 1998; Seemüller et al., 1998;

Adamovic et al., 2014PaeoniaceaePaeonia suffruticosa Tree peony Tree peony yellows 16SrXII China Gao et al., 2013



PapaveraceaeEscholtzia californica California poppy California poppy

yellows16SrI-A Canada Wang and Hiruki, 2005

Papaver rhoeas Corn poppy Papaver virescence 16SrI-B Italy Marcone et al., 2000

PedaliaceaePedalium murex Large caltrops Pedalium phyllody 16SrII-A India Babu et al., 2015Sesamum indicum Sesame Sesame phyllody 16SrI-B, 16SrII-A, -C, -D,

16SrIXAsia, Turkey Khan et al., 2007b; Win et al., 2010; Catal

et al., 2013; Ikten et al., 2014; Madhupriya et al., 2015b; Nabi et al., 2015, Singh et al., 2016

PhyllanthaceaePhyllanthus niruri Gale of the wind Phyllanthus little leaf 16SrI India Chaube et al., 2015b

PiperaceaePiper nigrum Black pepper Black pepper

phyllody16SrI India Bhat et al., 2006

Piper nigrum Black pepper Coorg black pepper yellows

16SrI-B India Adkar-Purushothama et al., 2009, 2011

PlantaginaceaePlantago lanceolata English plantain Plantain yellows 16SrI-B, 16SrXII-A Czech Republic, Italy Fránová and Šimková, 2009; Marcone,

2011; Mori et al., 2015Plantago major Broadleaf plantain Plantain yellows 16SrI-B, 16SrXII-A Germany, Italy, Serbia,

USAMarcone et al., 2000; Lee et al., 2004; Bellardi and Bertaccini, 2005; Borth et al., 2006; Josic et al., 2012; Mori et al., 2015

Plantago coronopus Plantain Plantain virescence 16SrI-A Germany Marcone et al., 2000; Lee et al., 2004

PlumbaginaceaeLimonium sinuatum Sea-lavender Limonium yellows 16SrI-A, -B, -C, -M Poland, Lithuania,

Canada, Iran, Japan, Korea

Kamińska et al., 1999; Valiunas et al., 2001; Lee et al., 2004; Chung et al., 2005; Samuitiene et al., 2007; Babaie et al., 2007; Maejima et al., 2014

Limonium tataricum German statice Limonium yellows 16SrI Canada Chang et al., 2004Limonium spp. Statice Limonium yellows 16SrI-B, 16SrII, 16SrV,

16SrIXItaly, Israel Gera et al., 2004; Abagnale et al., 2009;

Paltrinieri et al., 2016

PoaceaeCymbopogon citratus Lemon grass Cymbopogon citratus

white leaf16SrII India Madhupriya et al., 2014

Cynodon dactylon Bermuda grass Bermuda grass white leaf

16SrXIV-A, -C (‘Ca. P. cynodontis’)

Asia, Europe, Turkey, Sudan, Australia, Cuba

Marcone et al., 2004b; Arocha et al., 2005; Rao et al., 2007; Salehi et al., 2009; Çağlar et al., 2013; Khanna et al., 2015; Mitrović et al., 2015

Cynodon dactylon Bermuda grass Bermuda grass white leaf

16SrXII-A Serbia Mitrović et al., 2012

Digitaria sanguinalis Crab grass Digitaria white leaf 16SrXIV-A Italy, India Rao et al., 2010; Marcone, 2011

PolygonaceaePolygonum aviculare Common knotgrass Polygonum yellows 6SrVII-A, 16SrXII-A Chile, Italy Berger et al., 2009; Longone et al., 2011

PortulacaceaePortulaca grandiflora Moss-rose purslane Portulaca little leaf 16SrVI India Ajayakumar et al., 2007; Samad et al.,

2008Portulaca oleracea Purslane Portulaca yellows 16SrI-B; 16SrXII-A Italy Marcone et al., 2000; Mori et al., 2015

PrimulaceaePrimula spp. Primrose Primula yellows and

virescence16SrI-A, -B, -L Europe Lee et al., 2004; Fránová et al., 2016

PunicaceaePunica granatum Pomegranate Pomegranate yellows 16SrI-A, 16SrXII-A Turkey Karimi et al, 2015; Gazel et al., 2016

RanunculaceaeAconitum napellus Aconite Aconitum

proliferation16SrI-A, -L Lithuania Valiunas et al., 2001; Lee et al., 2004;

Samuitiene et al., 2007

RhamnaceaeHovenia dulcis Japanese raisin Japanese raisin

witches’ broom16SrV-B Asia Kamala-Kannan et al., 2011

Rhamnus catharticus Buckthorn Buckthorn witches’ broom

16SrXX-A (‘Ca. P. rhamni’) Europe Marcone et al., 2004a; Marcone, 2015

RosaceaeCrataegus monogyna Hawthorn Hawthorn yellows 16SrX-A, -C, -F Europe Tedeschi et al., 2009; Seemüller et al.,

2011Prunus spinosa Blackthorn European stone fruit

yellows16SrX-F Europe Marcone et al., 2010, 2011

Rosa alba Rose Rose little leaf 16SrI India Chaturvedi et al., 2009Rosa damascena Damask rose Rose witches’ broom 16SrV-B-related India Saeed et al., 2016Rosa multiflora Rose Rose dieback 16SrI-B Poland Kamińska et al., 2003



Rosa spp. Rose Rose yellows 16SrI, 16SrI-B, 16SrII, 16SrV-E, 16SrX-A, -C, -F

Iran, India, Italy, France, Poland

Jarausch et al., 2001; Kamińska et al., 2001; Kamińska and Śliwa, 2004; Bellardi and Bertaccini, 2005; Gao et al., 2008; Raj et al., 2011b; Aldaghi and Bertaccini, 2015

Rubus spp. Brambles Rubus stunt 16SrV-E (‘Ca. P. rubi’), 16SrI, 16SrIII, 16SrXII

Europe, Turkey, USA, Pakistan, New Zealand

Bertaccini et al., 1995a; Seemüller et al., 1998; Davies, 2000; Davis et al., 2001; Sertkaya et al., 2004; Borroto Fernández et al., 2007; Fahmeed et al., 2009; Reeder et al., 2010; Malembic-Maher et al., 2011; Cieslinska, 2011; Liefting et al., 2011

Spiraea salicifolia Bridewort Spiraea yellows 16SrV-B China Li et al., 2010bSpiraea spp. Spirea Spirea stunt 16SrIII-E USA Lee et al., 2000

RubiaceaeCoffea arabica Coffee Coffee crispiness 16SrIII Colombia Galvis et al., 2007Coffea arabica Coffee Coffee witches’

broom16SrIX Puerto Rico Caicedo et al., 2015

Ixora coccinea Ixora Ixora witches’ broom 16SrIX Puerto Rico Caicedo et al., 2015Morinda citrifolia Bartondi Bartondi yellows N.D. India Sarwade et al., 2015Rubia tinctorum Common madder Common madder

little leaf16SrVI-D Iran Zibadoost and Rastgou, 2016

RutaceaeCitrus paradisi Grapefruit Grapefruit yellows 16SrI, 16SrII-A-related China, Pakistan Fahmeed et al., 2009; Lou et al., 2014Dictamus alba Dittany Dictamus albus

yellows16SrI-B, 16SrIII-F Canada, Lithuania Wang and Hiruki, 2005; Valiunas et al.,

2007Zanthoxylum schinifolium Prickly ash Prickly ash witches’

broom16SrI Korea Han et al., 2013

SalicaceaeSalix alba White willow Salix witches’ broom 16SrXII-A Serbia Pavlović et al., 2014aSalix tetradenia Black mountain

willowSalix witches’ broom 16SrI China Mou et al., 2014

SantalaceaeSantalum album Sandal Sandal spike 16SrI-B India Lee et al., 2004; Khan et al., 2008

SapindaceaeDimocarpus longan Longan Longan witches’

broom16SrII, 16SrV, 16SrXII Vietnam Nguyen et al., 2012; Hoat et al., 2015

Melicoccus bijugatus Spanish lime Spanish lime fasciation

16SrIX Puerto Rico Caicedo et al., 2015

SapotaceaeManilkara zapota Sapodilla Sapodilla witches’

broom16SrII Cuba Acosta et al., 2009

ScrophulariaceaeDigitalis lanata Woolly foxglove Foxglove yellows 16SrI-B Italy Bellardi et al., 2007Digitalis lutea Yellow foxglove Yellow fowglow

yellows16SrI-B Italy Bellardi and Bertaccini, 2005

Digitalis purpurea Foxglove Foxglove yellows 16SrXII-A Serbia Pavlović et al., 2014aRehmannia glutinosa Chinese foxglove Rehmannia

proliferation16SrI-B Czech Republic Přibylová et al., 2001

SolanaceaeCapsicum annuum Chili Chili yellows 16SrI, 16SrII India, Indonesia Khan and Raj, 2006; Harling et al., 2009Cyphomandra betaceum Tamarillo Tamarillo yellows 16SrII Indonesia Harling et al., 2009Datura inoxia Downy thorn-apple Datura little leaf 16SrVI India Raj et al., 2009bDatura stramonium Common thorn-

appleDatura witches’ broom

16SrVI, 16SrXII-A India, Czech Republic, Greece, Turkey

Fialová et al., 2009; Özdemir et al., 2009; Lotos et al., 2013; Singh et al., 2013a, 2013b; Singh and Upadhyaya, 2013, 2015

Nicotiana tabacum Tobacco Tobacco “stolbur” 16SrXII-A Europe Seemüller et al., 1998; Marcone, 2011; Mitrović et al., 2013

Nicotiana tabacum Tobacco Tobacco big bud 16SrII Australia Schneider et al., 1999Solanum malacoxylon Waxyleaf

nightshadeWaxyleaf nightshade “stolbur”

16SrXII-A Italy Iriti et al., 2008

Solanum nigrum Black nightshade Unnamed 16SrII-E, 16SrIX-C Europe, Lebanon Battle et al., 2000; Tolu et al., 2006; Casati et al., 2016

Withania somnifera Indian ginseng Withania witches’ broom

16SrVI, 16SrI India Zaim and Samad, 1995; Khan et al., 2006; Samad et al., 2006

TamaricaceaeTamarix chinensis Salt cedar Salt cedar witches’

broom16SrXXX (‘Ca. P. tamaricis’)

China Zhao et al., 2005, 2009

TheaceaeCamellia japonica Camellia Camellia yellows 16SrV-B China Gao et al., 2015b



TiliaceaeTilia platyphyllos var. laciniata

Large-leaved linden Linden laciniated leaf 16SrI-B Lithuania Jomantiene et al., 2013

Tilia platyphyllos var. laciniata

Large-leaved linden Linden laciniated leaf witches’ broom

16SrI-(L/L)V Lithuania Jomantiene et al., 2013

TurneraceaeTurnera ulmifolia Yellow alder Turnera witches’

broom16SrXIII Brazil Montano et al., 2011b

UlmaceaeUlmus minor (syn. Ulmus campestris)

European field elm Elm yellows 16SrV-A (‘Ca. P. ulmi’) Europe, North America Marcone, 2015

UrticaceaeUrtica dioica Nettle Nettle yellows 16SrXII-A, 16SrII Europe, Israel Langer and Maixner, 2004; Bellardi and

Bertaccini, 2005; Aryan et al., 2014; Dafny Yelin et al., 2015; Mori et al., 2015; Sabaté et al., 2015; Kosovac et al., 2016

Parietaria officinalis Upright pellitory Pellitory yellows 16SrXII-A, 16SrX Italy Bellardi et al., 2004, 2008; Bellardi and Bertaccini, 2005

Parietaria judaica Spreading pellitory Pellitory yellows 16SrXII-A, 16SrX Italy Bellardi et al., 2004, 2008; Bellardi and Bertaccini, 2005

ValerianaceaeValeriana officinale Valerian Valerian yellows 16SrI-A, M, 16SrXII-A Canada, Lithuania,

SerbiaHwang et al., 1977b; Khadhair et al., 2008; Lee et al., 2004; Mitrović et al., 2013

VerbenaceaeVerbena x hybrida Vervain Verbena yellows 16SrI-B Turkey Přibylová et al., 2015Vitex agnus-castus Chaste tree Unnamed 16SrII Europe, Israel Dafny Yelin et al., 2015; Kosovac et al.,

2016

VitaceaeAmpelopsis brevipedunculata

Porcelain vine Porcelain vine witches’ broom

16SrI Korea Jung et al., 2003

Vitis vinifera Grapevine Grapevine yellows 16SrXII-A (‘Ca. P. solani’); 16SrXII-B (‘Ca. P. astraliense’); 16SrV-C, -D; 16SrVII-A (‘Ca. P. fraxini’-related); 16SrIX-C; 16SrX-B; 16SrIII-A (‘Ca. P. pruni’-related); 16SrIII-I, -J; 16SrI-A, -B, -C; 16SrII-D

Europe, USA, Australia, Israel, Chile, Tunisia, Iran, South Africa

Bertaccini et al., 1995b; Padovan et al., 1995; Davis et al., 1997b, 2015; M’hirsi et al., 2004; Belli et al., 2010; Bertaccini et al., 2014; Salehi et al., 2016b

a 16Sr group and subgroup designation according to Davis et al. (2016) and Zhao and Davis (2016), based on computer-simulated RFLP analysis. N.D., not determined.

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PHYTOPLASMA DISEASES OF MEDICINAL AND AROMATIC PLANTS