Stanley Freeman

• PhD
• Senior Researcher at Agricultural Research Organization ARO

Mango is affected by several fungal diseases, including dieback caused by species of the Botryosphaeriaceae family. Recently, mango dieback was reported from multiple locations in Israel. In this study, we isolated and characterized 11 representative fungal isolates belonging to Botryosphaeriaceae from symptomatic mango stems. Lasiodiplodia theobro...

In the present study, we investigated the in-vitro and in-vivo effects of root endophytic fungal species isolated from squirting cucumber Ecballium elaterium on morphological, physiological and biochemical mechanisms of muskmelon (Cucumis melo) in response to salt stress. A total of 23 endophytic fungal isolates belonging to nine different genera w...

Colletotrichum is an important phytopathogenic fungus that causes anthracnose disease in diverse agronomically important tropical food crops. Accurate pathogen identification is critical for early diagnosis and efficient management of anthracnose. ITS is not a reliable marker for this fungal genus due to its failure to phylogenetically resolve cryp...

Members of the Fusarium oxysporum complex are ubiquitous soilborne fungal pathogens causing wilt diseases in various plant hosts. Fusarium oxysporum (Fo) f. sp. cannabis was first reported causing wilt disease in hemp in Italy in 1962. To date, Fusarium wilt continues to cause concern in industrial and medicinal cannabis cultivation worldwide. Duri...

Fungi associated with cypress bark beetles are practically unknown in the Eastern Mediterranean. Our study focused on the fungi associated with the body parts and galleries of two indigenous cypress bark beetles, Phloeosinus armatus and P. bicolor, sampled from Cupressus sempervirens trees in different regions in Israel. Arbitrarily primed PCR, per...

Macrophomina phaseolina is a soil-borne fungal pathogen that incites charcoal rot in more than 500 plant species including melon, Cucumis melo. Disease incidence and severity are affected by host genetic background, plant age, temperature, and water economy. The non-genetic variation in disease severity exhibited by infected melon plants poses a ch...

Macrophomina phaseolina is a soil-borne fungal pathogen infecting many important crop plants. The fungus, which can survive on crop debris for a long period of time, causes charcoal rot disease by secreting a diverse array of cell-wall degrading enzymes and toxins. M. phaseolina thrives during periods of high temperatures and arid conditions, as ty...

Anthracnose caused by Colletotrichum spp. is an important disease of pepper (Capsicum annuum), affecting fruits and the overall yield worldwide. The disease was recently observed for the first time in Israel in the summer of 2020. In this study, morphology, phylogenetic analysis using glutamine synthase (gs) and the intergenic region between apn2 a...

The Ambrosia Fusarium Clade (AFC) is a monophyletic lineage within clade 3 of the Fusarium solani species complex (FSSC) that currently comprises 19 genealogically exclusive species. These fungi are known or predicted to be farmed by adult female Euwallacea ambrosia beetles as a nutritional mutualism (Coleoptera: Scolytinae; Xyleborini). To date, o...

Exotic diseases and pests of trees have caused continental-scale disturbances in forest ecosystems and industries, and their invasions are considered largely unpredictable. We tested the concept of pre-invasion assessment of not-yet invasive organisms, which enables empirical risk assessment of potential invasion and impact. Our example assesses fu...

Medical cannabis (MC) production is a rapidly expanding industry. Over the past ten years, many additional phytocannabinoids have been discovered and used for different purposes. MC was reported beneficial for the treatment of a variety of clinical conditions such as analgesia, multiple sclerosis, spinal cord injuries, Tourette's syndrome, epilepsy...

This study examined the polyphagous shot hole borer (PSHB) Euwallacea fornicatus (Coleoptera; Scolytinae) native to Southeast Asia and concentrated on its wide host range in two of the invaded areas, California and Israel. Among the 583 examined tree species, 55.9% were characterized as “non-reproductive hosts” and only 13.8% were characterized as...

Fusarium wilt, caused by Fusarium oxysporum is a major disease of jojoba, causing serious economic losses. This study aimed at characterizing the Fusarium populations associated with jojoba in Israel. Fifty Fusarium isolates used in this study included 23 isolates from the 90’s (‘past’) and 27 recently isolated (‘recent’). All the isolates were cha...

Background: The use of medical cannabis (MC) in the medical field has been expanding over the last decade, as more therapeutic beneficial properties of MC are discovered, ranging from general analgesics to anti-inflammatory and anti-bacterial treatments. Together with the intensified utilization of MC, concerns regarding the safety of usage, espec...

Scientific communication is facilitated by a data-driven, scientifically sound taxonomy that considers the end-user's needs and established successful practice. Previously (Geiser et al. 2013; Phytopathology 103:400-408. 2013), the Fusarium community voiced near unanimous support for a concept of Fusarium that represented a clade comprising all agr...

The use of and research on medical cannabis (MC) is becoming more common, yet there are still many challenges regarding plant diseases of this crop. For example, there is a lack of formal and professional knowledge regarding fungi that infect MC plants, and practical and effective methods for managing the casual agents of disease are limited. The p...

Macrophomina phaseolina is a typical soilborne fungal pathogen causing crown and root rot in strawberry (Fragaria × ananassa Duch.) worldwide. M. phaseolina has become a major problem for strawberry growers parallel to the phase-out of methyl-bromide since 2004 and is considered the most destructive soilborne pathogen of strawberry since then. Glob...

Fusarium oxysporum f.sp. cubense (Foc) is considered one of the most devastating soilborne fungal pathogens of banana worldwide. Foc causing mortality to Cavendish group bananas, and belonging to the unique vegetative compatibility group (VCG) 01213/16 has been termed tropical race 4 (TR4) and has currently been renamed F. odoratissimum. The pathog...

The Ambrosia Fusarium Clade (AFC) comprises at least 16 genealogically exclusive species-level lineages within clade 3 of the Fusarium solani species complex (FSSC). These fungi are either known or predicted to be farmed by Asian Euwallacea ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) in the tribe Xyleborini as a source of nutrition. To...

Interactions between the fungal symbionts of the polyphagous shot hole borer (PSHB) Euwallacea nr. fornicatus and avocado host trees of cultivars 'Hass' and 'Ettinger' were studied, with emphasis on the three symbiotic fungi of the beetle. Fusarium euwallaceae did not spread far from the beetle galleries and remained viable in live xylem for 25 mon...

It has long been known that hormones affect the interaction of a phytopathogen with its host plant. The pathogen can cause changes in plant hormone homeostasis directly by affecting biosynthesis or metabolism in the plant or by synthesizing and secreting the hormone itself. We previously demonstrated that pathogenic fungi of the Fusarium species co...

Anthracnose disease caused by Colletotrichum species is a major constraint for the shelf-life and marketability of avocado fruits. To date, only C. gloeosporioides sensu lato and C. aenigma have been reported as pathogens affecting avocado in Israel. This study was conducted to identify and characterize Colletotrichum species associated with avocad...

The polyphagous shot hole borer (PSHB), Euwallacea nr. fornicatus (Coleoptera; Scolytinae) has become a serious threat to the avocado industry and several shade tree species in Israel. Branch wilting and tree mortality is the outcome of PSHB galleries. Understanding the relationship between avocado trees and the PSHB is required for considering man...

Species of the Fusarium fujikuroi species complex (FFC) cause a wide spectrum of often devastating diseases on diverse agricultural crops, including coffee, fig, mango, maize, rice, and sugarcane. Although species within the FFC are difficult to distinguish by morphology, and their genes often share 90% sequence similarity, they can differ in host...

Mango malformation disease (MMD), caused by Fusarium mangiferae, is a major constraint to mango production, causing significant yield reduction resulting in severe economic impact. The present study characterizes fungal localisation in planta during initiation and development of vegetative and floral malformation. Young mango trees were artificiall...

Several species of the ambrosia beetle Euwallacea (Coleoptera: Curculionidae: Scolytinae) cultivate Ambrosia Fusarium Clade (AFC) species in their galleries as a source of food. Like all other scolytine beetles in the tribe Xyleborini, Euwallacea are thought to be obligate mutualists with their fungal symbionts. Published diversification-time estim...

The Asian ambrosia beetle, Euwallacea nr. fornicatus carries three species of symbiotic fungi namely, Fusarium euwallaceae, Graphium euwallaceae and Paracremonium pembeum; all three fungi are cultivated within the galleries excavated in the host tree xylem. Among the 50 tree species attacked by the beetle few ornamental and several avocado varietie...

The presence of three symbiotic fungi: Fusarium euwallaceae, Graphium euwallaceae and Acremonium pembeum was detected in the larvae and adult beetles of Euwallacea nr. fornicatus (Coleoptera: Scolytinae) and from the brood galleries of four tree species. Isolations from female beetle mandibular mycangia recovered mainly G. euwallaceae and A. pembeu...

Avocado (Persea americana) is widely cultivated throughout Israel, with the coastal plain and eastern valleys of Upper Galilee comprising the major growing areas (Dor 2015). During sample collection of unripe avocado fruits (cv. Hass) from northern and central Israel in November 2014, profuse salmon-colored sporulation was observed on the skins of...

Diagnosis and diversity of the Fusarium spp. isolates causing mango malformation in Southern Spain by using molecular and genetic techniques.

Mango malformation disease (MMD) has become an important global disease affecting this crop. The aim of this study was to identify the main causal agents of MMD in the Axarquia region of southern Spain and determine their genetic diversity. Fusarium nzangiferae was previously described in the Axarquia region but it represented only one-third of the...

Fungi associated with the bark beetles Orthotomicus erosus, Tomicus destruens and Pityogenes calcaratus were sampled in various pine forests throughout Israel. Three ophiostomatoid fungi, Ophiostoma ips, Graphilbum rectangulosporium and Leptographium wingfieldii, and a fourth non-ophiostomatoid fungus, Geosmithia sp. 24, were identified by using mo...

Panama disease (aka Fusarium wilt) of banana (Musa spp.) has been a destructive problem for well over a century. Race 1 of the pathogen, Fusarium oxysporum f. sp. cubense (Foc), was responsible for the demise of the first export trades of banana that were based on the cultivar ‘Gros Michel’. Currently, tropical race 4 (TR4) impacts the Cavendish cu...

Malformation is the most important disease of mango in Mexico. It affects floral and vegetative shoots, and reduces fruit production. Although several species of Fusarium cause the disease worldwide, F. mexicanum is most important in Mexico. To better understand epidemiology of disease in Mexico, we studied mango nurseries as a source of infected p...

Mango malformation disease (MMD) caused by Fusarium mangiferae severely affects the crop and is widely distributed in almost all mango-growing regions worldwide. Additional Fusarium species have also been reported to cause MMD. Since malformed inflorescences do not bear fruit, MMD is a major constraint to crop production in affected areas. Symptoms...

Mango (Mangifera indica L.) malformation disease (MMD) is one of the most important diseases affecting this crop worldwide, causing severe economic loss due to reduction of yield. After the first report in India in 1891 (3), MMD has spread worldwide to most mango-growing regions. Several species of Fusarium cause the disease, including F. mangifera...

Mango (Mangifera indica) is regarded as the king of fruits in India, where it has been cultivated for at least 4,000 years and has great cultural and religious significance. Many Indian mango cultivars originated in the fifteenth century when the best selections of mango seedlings were propagated by grafting and planted in large orchards, in some c...

In the past 10 years, there has been a substantial increase in reports, from growers and extension personnel, on bulb and root rots in lily (Lilium longiflorum) in Israel. Rot in these plants, when grown as cut flowers, caused serious economic damage expressed in reduction in yield and quality. In lily, the fungal pathogens involved in the rot were...

Fusarium root and stem rot caused by the fungus Fusarium oxysporum f. sp. radicis-cucumerinum is a major disease in greenhouse cucumbers. Over the past decade, the disease has been documented in melon greenhouses in Greece, and recently it has been sporadically recorded in greenhouse melons in Israel. Variations in disease response were found among...

Mango malformation disease (MMD) caused by Fusarium mangiferae severely affects the crop and is widely distributed in almost all mango-growing regions worldwide. Since malformed inflorescences do not bear fruit, MMD is a major constraint to crop production in affected areas. No effective management methods have been reported to date. The airborne n...

During the last years, Fusarium strains have been isolated from shoots and inflorescences of mango trees affected with floral and vegetative malformation in different orchards from the Axarquia region (south of Spain), highlighting the identification of Fusarium mangiferae. With the aim of elucidate epidemiological aspects and design more efficient...

During the last years, Fusarium strains have been isolated from shoots and inflorescences of mango trees affected with floral and vegetative malformation in different orchards from the Axarquía region (south of Spain), highlighting the identification of Fusarium mangiferae. With the aim of elucidate epidemiological aspects and design more efficient...

The ambrosia beetle, Euwallacea aff. fornicata Eichhoff was first recorded in Israel in 2009. The symbiotic fungus Fusarium sp. nov., carried in the mandibular mycangia of the beetle, is responsible for typical wilt and dieback symptoms inflicted on avocado (Persea americana Miller) trees. The beetle-fungus complex has become a serious threat to th...

The invasive Asian ambrosia beetle Euwallacea sp. (Coleoptera, Scolytinae, Xyleborini) and a novel Fusarium sp. that it farms in its galleries as a source of nutrition causes serious damage to more than 20 species of live trees and pose a serious threat to avocado production (Persea americana) in Israel and California. Adult female beetles are equi...

Variability of chromosomes XII and IV. A: PCR analysis reveals that chromosome XII is not present in all F. fujikuroi strains. 1, 2 and 3: PCR products derived from primer pairs designed from the sequence of genes FFUJ_14099, FFUJ_ 14193, and FFUJ_ 14245, respectively. The genes FFUJ_14099, FFUJ_14193, and FFUJ_14245 are located 0.7, 0.4 and 0.08 M...

Maximum likelihood trees of selected GA biosynthetic genes. A: tree generated from Cps/Ks and related diterpene synthases from multiple genera of fungi; B: tree generated from concatenated sequences of GA enzymes/genes common to Fusarium species, Claviceps purpurea and Sphaceloma manihoticola. C: trees of four GA biosynthetic enzymes/genes (P450-1,...

Phylogenetic tree of PKSs encoded by Fusarium genome sequences. The tree was generated by maximum parsimony analysis of alignments of deduced amino acid sequences of the KS and AT/MAT domains. The tree also includes PKSs from other fungi and for which the corresponding secondary metabolites is known. The names of metabolites are indicated to the ri...

Comparison of the bikaverin biosynthetic (BIK) cluster in genomes sequences of Fusarium. BIK genes are represented by blue horizontal arrows, and the numbers within these arrows correspond to BIK gene numbers rather than gene/protein model designations from genome sequence databases. For those that are available, gene designations are indicated bel...

Comparison of the putative, two-gene PKS12 clusters (green arrows) in genome sequences of Fusarium. The second gene in the cluster is predicted to encode a methyltransferase. Blue arrows represent cluster flanking genes, which exhibit partial synteny conservation among the species examined. Based on phylogenetic analysis (Figure 5), PKS12 homologue...

Expression pattern and distribution of active histone marks at the A: Fusaric acid (FUB) and B: at the fumonisin gene cluster. (TIF)

CHEF gel analysis of F. fujikuroi IMI58289. Previously we performed Southern blot analyses [39] of the CHEF gel with the following gene probes: CREA (FFUJ_04790), hmgR (FFUJ_04000), NMR (FFUJ_02636), GGS1(FFUJ_07352), CPS/KS (FFUJ_1433), AREA (FFUJ_06143), NIAD (FFUJ_12277), CARRA (FFUJ_11802). Arrows show the position of the hybridization signals,...

Characterization of F. fujikuroi chromosomes I–XII: variation in GC-content, acetylation and methylation statues of histone H3 and expression under low nitrogen conditions. For each chromosome, a diagram showing the position of the centromere is shown at the top; below this in descending order are: GC content, location of SM biosynthetic gene clust...

Relative expression of the fumonisin biosynthetic gene FUM1 from F. fujikuroi and F. verticillioides in rice and maize roots. Rice (A) and maize (B) roots were infected with Fusarium fujikuroi and F. verticillioides spores and every 2 days RNA was isolated from three or five plants and used in real time PCR analysis. The expression levels were obta...

Putative F. fujikuroi centromeric regions. Coordinates on chromosomes (Chr) I to XII include 0.5–1 kb of euchromatic regions on either side of pericentric and centromeric DNA and no attempt has been made to separate pericentric and centromeric regions. As none of the 14 scaffolds contain telomere repeats, all chromosome ends are unfinished. “Incomp...

Gibberellin biosynthesis in species of the GFC. A: Expression of gibberellins biosynthetic genes cps/ks and des after three days of cultivation under nitrogen-limiting conditions (6 mM glutamine). All strains except for F. verticillioides contain the entire GA gene cluster. B: GC-MS analysis of ent-kaurene, the first specific intermediate of the GA...

Comparison of the fusarubin biosynthetic gene (FSR/PGL) cluster in genomes sequences of Fusarium. Horizontal arrows that are the same color represent genes, or gene sets, that have closely related homologues in two or more species. Blue arrows represent FSR/PGL genes, and the numbers within these arrows correspond to FSR/PGL gene numbers rather tha...

Comparison of the putative fusaric acid biosynthetic gene (FUB) cluster in genomes sequences of Fusarium. Horizontal arrows that are the same color represent genes, or gene sets, that have closely related homologues in two or more species. Blue arrows represent FUB genes, and the numbers within these arrows correspond to FUB gene numbers rather tha...

Comparison of the putative PKS17 – PKS18 gene cluster in the genome sequences of F. fujikuroi and F. mangiferae. The cluster genes are represented by green horizontal arrows, and cluster flanking genes are represented by blue or yellow arrows; yellow arrows represent genes that were not detected in the flanking regions of F. fujikuroi or F. mangife...

Identification of secondary metabolite gene clusters based on SMURF and sequence comparisons with related fungal genomes. (XLSX)

Comparison of the fusarin biosynthetic gene (FUS) cluster in genome sequences of Fusarium and the related fungus Metarhizium anisopliae as well as remnants of the FUS cluster in F. mangiferae and Trichoderma reesei. FUS genes are represented by blue horizontal arrows, and the numbers in the arrows correspond to FUS gene numbers. For those that are...

Phylogenetic tree of Fusarium NRPS22 and Beauveria bassiana BeaS and BslS as well as F. scirpi EsyN. The tree was generated by maximum parsimony analysis of alignements of deduced amino acid sequences of the A domains. The protein/gene designations FCIRG, FFUJ, FOXG and FVEG correspond to NRPSs deduced from genome sequences of F. circinatum, F. fuj...

Venn diagram representing the distribution of nitrogen-and pH-regulated genes. Number of differentially regulated genes in wild-type F. fujikuroi cultivated under conditions that vary in nitrogen content and pH. The data were obtained by microarray analyses. The conditions were as follows: 6 mM gln (glutamine) constitutes, acidic low nitrogen; 60 m...

Nearest genes to F. fujikuroi centromeres and synteny with F. verticillioides. Nearest F. fujikuroi genes (FFUJ locus ID) to the left and right of the predicted centromere on each chromosome was identified and compared to the location of predicted orthologs in F. verticillioides (FVEG locus ID). “+” and “−” indicate strands, “t” indicates last gene...

Overview of the analysed amino acids. The amino acids that are obtained after acid hydrolysis of apicidin and the apicidin metabolite were identified based on the accurate mass of their [M+H]+ ions. (DOCX)

Gene family features, comparative to related species. (XLSX)

Gibberellin production by Fusarium spp. A: Gibberellin production by Fusarium species of the GFC and F. oxysporum 4287 (outgroup). F. oxysporum 4287 does not contain a GA gene cluster, but produces GAs after transforming it with the entire GA gene cluster from F. fujikuroi (cos1). B: Gibberellin production by F. oxysporum isolates with an entire GA...

Secondary metabolite production in F. fujikuroi IMI58289, F. circinatum Fsp34, F. mangiferae MRC7560, F. verticillioides 3125 and F. oxysporum 4287. The strains were cultivated under four different culture conditions. Bikaverin, O-methylfusarubin, fusarins, fumonisin B1, fusaric acid and beauvericin were analyzed by HPLC-FTMS. (DOCX)

Primers used for all experiments. (DOCX)

Expression pattern of all genes under four standard growth conditions. acidic and alkaline low-nitrogen (6 mM glutamine or 6 mM sodium nitrate) and acidic and alkaline high-nitrogen (60 mM glutamine or 120 mM sodium nitrate), as well as proteomic data and the presence of activation histone marks under two different growth conditions (acidic low and...

Overview of the metabolites that were analyzed by HPLC-FTMS. The analytes were identified by their retention time and isotope pattern compared to standard substances. For the estimation of the produced amount, the m/z of their most intense ions were observed. (DOCX)

The fungus Fusarium fujikuroi causes "bakanae" disease of rice due to its ability to produce gibberellins (GAs), but it is also known for producing harmful mycotoxins. However, the genetic capacity for the whole arsenal of natural compounds and their role in the fungus' interaction with rice remained unknown. Here, we present a high-quality genome...

The following is a summary of Fusaria associated with mango malformation disease (MMD) in México. From 2002 to 2009, 142 strains were isolated from symptomatic mango inflorescences and vegetative tissues from various cultivars in eight geographically diverse Mexican states. Initially, isolates were screened for genetic diversity using arbitrarily p...

Mango malformation disease (MMD) is one of the most devastating diseases causing severe economic losses to this crop worldwide. MMD has not been reported in Sri Lanka although the disease was reported in neighboring India over a century ago. Abnormal, thick, and fleshy mango panicles (40%) and proliferating stunted shoots (<1%) showing characterist...

In this letter, we advocate recognizing the genus Fusarium as the sole name for a group that includes virtually all Fusarium species of importance in plant pathology, mycotoxicology, medicine and basic research. This phylogenetically-guided circumscription will free scientists from any obligation to use other genus names, including teleomorphs, for...

Colletotrichum and its teleomorph Glomerella are considered major fungal plant pathogens worldwide. They cause significant economic damage to fruit crops in tropical, subtropical and temperate regions. Several Colletotrichum species or biotypes are known to cause disease in a single host such as C. acutatum and C. gloeosporioides on apple, avocado,...

The ambrosia beetle, Euwallacea fornicatus Eichhoff, was first recorded in Israel in 2009. The symbiotic fungus Fusarium sp., carried in mycacangia located in the anterior region of the female head, is responsible for the typical wilt symptoms inflicted on avocado (Persea americana Miller) trees. The beetle–fungus complex has become a serious threa...

Mango malformation disease (MMD) caused by various Fusaria, including the fungus Fusarium mangiferae, is difficult to diagnose and cannot be controlled effectively. In this study a PCR diagnostic tool was developed for detection of F. mangiferae by generating primers which flank fungus-specific sequences. The assay is suitable for both in-vitro and...

Identificator is a web-based tool used to help non experts in identifying plant diseases, based on the selection of pictures and/or short text descriptions (when no suitable images exist) representing the symptoms on a specific sample of plant organs. The system is based on a multi-access key of identification and specifically on the selection of p...

Mango (Mangifera indica L.) malformation disease (MMD) is one of the most important diseases affecting this crop worldwide, which causes severe economic losses because of the reduction of productivity. Symptoms of MMD in Spain were observed for the first time in April of 2006 in three mango orchards in the Axarquia Region (southern Spain). Symptoms...

Zveibil, A., Mor, N., Gnayem, N., and Freeman, S. 2012. Survival, host pathogen interaction, and management of Macrophomina phaseolina on strawberry in Israel. Plant Dis. 96:265-272. Crown and root rot of strawberry, caused by Macrophomina phaseolina, have become predominant soilborne diseases of strawberry in Israel over the past 5 years. In total...

Anthracnose of strawberry caused by Colletotrichum acutatum was recently recorded in Egypt in 2006, however, reliable molecular identification was not conducted at that time. During 2007 and 2008, typical anthracnose symptoms were observed in various cultivars of field-grown strawberry in Qalubia and Ismailia governorates. Lesions on necrotic petio...

La malformacion del mango es una de las enfermedads más importantes que afectan a este cultivo en todo el mundo, causando imporantes perdidas economicas. Este trabajo pone de manifiesto de manera concluyente la presencia por vez primera de la malformación del mango en España y la determinacion de su agente causal