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Pines are the most economically important trees in the world and, together with eucalyptus, they dominate commercial forests. But the success of a relatively small number of widely planted species, such as Pinus pinaster, the maritime pine, comes at a price. Pines are attractive to damaging pathogens and insect pests, including the pinewood nematode (PWN), Bursaphelenchus xylophilus, the causal agent of pine wilt disease (PWD). Originally described in Japan, PWD has caused widespread destruction to forests in countries such as China, Taiwan, Portugal, Spain and the United States. PWN causes irreparable damage to the vascular system of its pine hosts, leading to mortality within 3 months. Pine sawyer beetles (Monochamus spp.) are key vectors of PWD, introducing the PWN to healthy trees during feeding. Other organisms contribute to PWD spread and development, including bacteria, fungi and bark beetles. Control measures include tree felling to prevent vector transmission of PWN, insecticide treatments, trapping of Monochamus spp. and tree breeding for plant resistance. The PWN is a quarantine pathogen and subject to regular legislation and phytosanitary measures aimed at restricting movement and preventing introduction to new areas. Current research is investigating the use of biopesticides against PWN and Monochamus spp. This review examines the biology, epidemiology, impact and management of PWD through published research, grey literature and interviews with people directly involved in the management of the disease in Portugal. [ABSTRACT FROM AUTHOR]

Control of the pinewood nematode (PWN), the causal agent of pine wilt disease, can be achieved through the trunk injection of nematicides; however, many pesticides have been linked to environmental and human health concerns. Essential oils (EOs) are suitable alternatives due to their biodegradability and low toxicity to mammals. These complex mixtures of plant volatiles often display multiple biological activities and synergistic interactions between their compounds. The present work profiled the toxicity of eight EOs against the PWN in comparison to their 1:1 mixtures, to screen for successful synergistic interactions. Additionally, the main compounds of the most synergistic mixtures were characterized for their predicted environmental fate and toxicity to mammals in comparison to emamectin benzoate, a commercial nematicide used against PWN. The mixtures of Cymbopogon citratus with Mentha piperita and of Foeniculum vulgare with Satureja montana EOs showed the highest activities, with half-maximal effective concentrations (EC50) of 0.09 and 0.05 µL/mL, respectively. For these, complete PWN mortality was reached after only ca. 15 min or 2 h of direct contact, respectively. Their major compounds had a higher predicted affinity to air and water environmental compartments and are reported to have very low toxicity to mammals, with low acute oral and dermal toxicities. In comparison, emamectin benzoate showed lower nematicidal activity, a higher affinity to the soil and sediments environmental compartments and higher reported oral and dermal toxicity to mammals. Overall, uncovering synergistic activities in combinations of EOs from plants of different families may prove to be a source of biopesticides with optimized toxicity against PWNs. [ABSTRACT FROM AUTHOR]

Considered one of the most devastating plant–parasitic nematodes worldwide, Bursaphelenchus xylophilus (commonly known as pinewood nematode, PWN) is the causal agent of the pine wilt disease in the Eurasian coniferous forests. This migratory parasitic nematode is carried by an insect vector (Monochamus spp.) into the host tree (Pinus species), where it can feed on parenchymal cells and reproduce massively, resulting in the tree wilting. In declining trees, PWN populations are strongly dependent on fungal communities colonizing the host (predominantly ophiostomatoid fungi known to cause sapwood blue-staining, the blue-stain fungi), which not only influence their development and life cycle but also the number of individuals carried by the insect vector into a new host. Our main aim is to understand if PWN-associated mycobiota plays a key role in the development of PWD, in interaction with the PWN and the insect vector, and to what extent it can be targeted to disrupt the disease cycle. For this purpose, we characterized the fungal communities of Pinus pinaster trees infected and non-infected with PWN in three collection sites in Continental Portugal with different PWD temporal incidences. Our results showed that non-infected P. pinaster mycoflora is more diverse (in terms of abundance and fungal richness) than PWN-infected pine trees in the most recent PWD foci, as opposed to the fungal communities of long-term PWD history sites. Then, due to their ecological importance for PWN survival, representatives of the main ophiostomatoid fungi isolated (Ophiostoma, Leptographium , and Graphilbum) were characterized for their adaptative response to temperature, competition in-between taxa, and as food source for PWN. Under the conditions studied, Leptographium isolates showed promising results for PWN control. They could outcompete the other species, especially O. ips , and significantly reduce the development of PWN populations when compared to Botrytis cinerea (routinely used for PWN lab culturing), suggesting this to be a natural antagonist not only for the other blue-stain species but also for the PWN. [ABSTRACT FROM AUTHOR]

Pine wilt disease (PWD), caused by the pinewood nematode (PWN) Bursaphelenchus xylophilus, is one of the most destructive diseases in trees of the genus Pinus and is responsible for significant environmental and economic losses in North America, Eastern Asia, and Western Europe. However, pine species are not equally affected, with some being tolerant/resistant while others are susceptible to nematode infection for reasons still unclear. The present study aims to investigate differential chemical responses of susceptible and tolerant/resistant pine species shortly after nematode infection by characterizing the phenolic profiles of adult Pinus sylvestris, Pinus pinaster, Pinus pinea, and Pinus halepensis. HPLC and LC-MS were used to identify and quantify the pine´s phenolic compounds: gallic acid, ferulic acid, taxifolin, rutin, resveratrol, (+)-secoisolariciresinol, (−)-epicatechin, protocatechuic acid hexoside, gallic acid hexoside, ferulic acid glucoside, quercetin hexoside, and two unidentified compounds (#A and #B). Prior to infection, we could not differentiate between nematode-tolerant/resistant and susceptible adult pine species based on their constitutive phenolic compounds. In the presence of the PWN, the phenolic profile allowed for a noticeable separation of the PWN-tolerant/resistant P. halepensis from the susceptible P. sylvestris, contrasting with a more homogenous response from P. pinea and P. pinaster. Observations on P. halepensis suggest that taxifolin, resveratrol, and rutin may have an active role in protecting against B. xylophilus, possibly in conjugation with other biochemical and anatomical characters. We emphasize the importance of studying pine tolerant/resistance on adult trees, and not on excised branches, saplings, or seedlings to accurately simulate the nematode–pine host interactions occurring under natural conditions. [ABSTRACT FROM AUTHOR]

Pine wilt disease (PWD) is a complex disease that severely affects the biodiversity and economy of Eurasian coniferous forests. Three factors are described as the main elements of the disease: the pinewood nematode (PWN) Bursaphelenchus xylophilus, the insect-vector Monochamus spp., and the host tree, mainly Pinus spp. Nonetheless, other microbial interactors have also been considered. The study of mycoflora in PWD dates back the late seventies. Culturomic studies have revealed diverse fungal communities associated with all PWD key players, composed frequently of saprophytic fungi (i.e., Aspergillus, Fusarium, Trichoderma) but also of necrotrophic pathogens associated with bark beetles, such as ophiostomatoid or blue-stain fungi. In particular, the ophiostomatoid fungi often recovered from wilted pine trees or insect pupal chambers/tunnels, are considered crucial for nematode multiplication and distribution in the host tree. Naturally occurring mycoflora, reported as possible biocontrol agents of the nematode, are also discussed in this review. This review discloses the contrasting effects of fungal communities in PWD and highlights promising fungal species as sources of PWD biocontrol in the framework of sustainable pest management actions. [ABSTRACT FROM AUTHOR]

Esteya vermicola is a potential biological agent of the pinewood nematode (PWN), Bursaphelenchus xylophilus, due to its high infectivity. However, knowledge of E. vermicola colonization in the host pine tree is less known. To reveal the distribution pattern of E. vermicola inside the pine tree and the interactions between fungus and PWN, biocontrol tests were conducted on 10 years old Pinus densiflora. A green fluorescence protein (GFP)‐tagged E. vermicola was used to observe the fungal hyphae. Real‐time TaqMan PCR quantification was applied to quantify the fungal hyphae in host tree. The results suggest that inoculation with the fungus significantly improved the survival rate of tested trees. Besides, the number of PWN extracted from fungal inoculated tested trees significantly decreased. Moreover, the fungal hyphae and lunate conidia grown in tested trees were observed with the GFP‐tagged E. vermicola. The real‐time quantification of E. vermicola illustrated the differential distribution of E. vermicola in wilted, wilting and healthy pine trees. Additionally, the promotion of fungal hyphal dispersion by B. xylophilus was found in the pine trees artificially infected with PWN. This study reveals the distribution of E. vermicola and PWN during biocontrol of pine wilt disease and provides direct evidence of the effectiveness of this fungus to control PWN. The migration of PWN infected by E. vermicola was supposed to improve the fungal extension in host pine tree. [ABSTRACT FROM AUTHOR]

The pinewood nematode (PWN), Bursaphelenchus xylophilus, is the causal agent of pine wilt disease (PWD) and a quarantine organism in many countries. Managing PWD involves strict regulations and heavy contingency plans, and present climate change scenarios predict a spread of the disease. The urgent need for sustainable management strategies has led to an increasing interest in promising biocontrol agents capable of suppressing the PWN, like endoparasitic nematophagous fungi of the Esteya genus. Here, we review different aspects of the biology and ecology of these nematophagous fungi and provide future prospects. [ABSTRACT FROM AUTHOR]