Plant microbiome research has been expanding over the last years due to its key determinant role in plant health and crop productivity [1], as is revealed by the increase in the number of scientific publications listed in Web of Science (WoS) and Scopus databases. Nowadays, it is generally accepted that plants live in association with a rich diversity of microorganisms. The sum of these microbial cells formed by multi-kingdom microbial communities (bacteria, fungi, viruses, etc.) that colonize both below- and aboveground plant organs in a particular environment is called plant-associated microbiota and refers to the taxonomy and abundance of microbial community members in a given environment. On the other hand, the totality of genomes of this microbiota is called the plant microbiome and it is often used to define the microbial behaviors or functions determined by a microbiota [2].

A better knowledge of the plant microbiome will result in developing alternatives to solve some of the environmental issues and challenges worldwide caused by excess use of chemicals for pest control by enhancing crop production and soil health [3] and by developing biological control alternatives for managing plant diseases ^[4][5][4,5]. Plant-associated beneficial microbiomes can confer fitness advantages to the host plant because these microorganisms can directly impact plant growth and health by producing phytohormones, improving nutrient acquisition and phosphate solubilization and tolerance to biotic and abiotic stresses. Additionally, they can act indirectly by activating a series of host physiological and immune responses that may trigger the multiplication of a specific microbiome to cope with the pathogen infection. This beneficial elicited microbiome can compete for plant resources and niche space or inhibit the growth of plant pathogens via the production of antibiotics, fungal cell wall-degrading enzymes or siderophores, among others ^[6][7][8][6,7,8].

Several authors have described the importance and ecological interpretations of the cross-kingdom co-occurrence networks in the plant microbiome ^[9][10][162,163]. Network analyses have identified the existence of interactions among the myriad of endophytes present in natural communities and other potential higher-order interactions ^[11][164]. Thus, this analysis has contributed to the discovery of hub or keystone microbial taxa in the plant microbiome. In fact, leaves of Arabidopsis thaliana, a fungal species, Dioszegia sp., and an oomycete, Albugo sp., were detected as hubs in the networks and experimentally validated as keystone species ^[12][165]; whereas in the wild rice seed microbiome, two fungal species were identified to be hub species in the cross-kingdom network ^[13][166]. This data analysis tool can be used to determine the existence of aggregation or exclusion interactions in xylem-associated microbiomes and reveal keystone microbial players in the xylem of olive trees to fight against vascular pathogens to contribute to the production of olive plants that are more resilient to these pathogens.

2. Olive Tree and Its Vascular Pathogens

Olive (Olea europaea susbp. europaea L.) is one of the most relevant tree species within the Mediterranean Basin due to its cultural and economic importance, without underestimating the noteworthy environmental attributes related to its ability to survive in poor, shallow and dry soils, and its contribution to the control of soil erosion and increase in soil nutrient retention ^[14][15][9,10]. In addition, olive orchards are important agroforestry habitats, playing essential roles in maintaining ecosystem diversity ^[16][11]. Hence, olive cultivation comprises relevant economic, environmental and socio-cultural values representing an agricultural system that should remain integral to the Mediterranean Basin. However, the health status of olive trees is being threatened by a remarkable increase in diseases caused by xylem-inhabiting pathogens, such as the plant pathogenic bacterium Xylella fastidiosa and the soilborne fungus Verticillium dahliae. Both pathogens are global threats to olive production which can adversely affect olive growth and production, causing substantial economic losses and severe environmental impact ^{[17][18][19][20][21]}[12,13,14,15,16]. The life cycle of V. dahliae, the causal agent of Verticillium wilt, is mainly characterized by its growth confined within the xylem vessels during the pathogenic phase and its ability to survive for many years as dormant microsclerotia in soil or within plant debris ^[17][22][12,17]. Verticillium dahliae infects the olive plants through the root system; then, the pathogen colonizes the xylem vessels and impairs the sap flow through mycelial proliferation, the formation of occlusions and tyloses that ultimately may cause the death of the tree ^[17][23][12,18]. The implementation of an integrated management strategy is the best option for controlling this disease, combining the use of resistant olive cultivars, or tolerant ones grafted onto resistant rootstocks, with adequate irrigation management and agricultural practices that prevent the spread of inoculum of the pathogen ^{[17][24][25][26][27][28]}[12,19,20,21,22,23]. On the other hand, X. fastidiosa is a quarantine plant pathogenic bacterium in the European Union (EU) and was ranked first on the list of the 20 priority pests within the EU territory (Commission Delegated Regulation (EU) 2019/1702). X. fastidiosa is transmitted by sap-feeding insect vectors and has been described as the causal agent of Olive Quick Decline Syndrome (OQDS). This disease is characterized by rapid dieback of shoots, twigs and branches, followed by the death of the tree ^[29][24]. Sap-feeding insect vectors inoculate the pathogen into xylem vessels from infected to healthy trees. Once inside, the aggregation of bacterial cells and biofilm formation block the sap flow in the xylem causing the plant to dry out and die ^[30][25]. Xylella fastidiosa is taxonomically divided into three major subspecies (fastidiosa, multiplex and pauca) ^[31][26], although two additional subspecies have been proposed (morus and sandyi). Only isolates belonging to the subspecies pauca are currently considered a threat to olive production worldwide. As previously indicated for V. dahliae, currently, there is no efficient control measure for the management of X. fastidiosa once the plant is infected. Consequently, the use of preventive strategies focused on the early detection and eradication of infected host plants, the control of the vectors and restrictions on plant material movements are currently the only available tools for the management of the diseases caused by this harmful bacterium ^[32][33][34][27,28,29]. The establishment of a pathogen lifestyle in a microorganism depends on its interactions with the host microbiome and the host immune system ^[35][30]. The xylem-associated microorganisms are involved in several biotic and abiotic processes within the plant host, including the acquisition of nutrients and increase of plant tolerance to abiotic stresses, without overlooking their role in the defense of the plant against vascular pathogens. In this context, acquiring and maintaining a beneficial xylem-associated microbiota capable of adapting more rapidly to a changing environment could be a selective advantage for the olive tree to fight against vascular pathogens ^[36][37][31,32].