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ah/pan/jam x sativa seeds

Ah/pan/jam x sativa seeds
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AH/PAN/JAM x Sativa

breed by B. Seeds Co.

Here you can find all info about AH/PAN/JAM x Sativa from B. Seeds Co.. If you are searching for information about AH/PAN/JAM x Sativa from B. Seeds Co., check out our Basic Infos, Shop-Finder and Price Comparison or Lineage / Genealogy for this cannabis variety here at this page and follow the links to get even more information. If you have any personal experiences with growing or consuming this cannabis variety, please use the upload links to add them to the database!

Basic / Breeders Info

AH/PAN/JAM x Sativa is a mostly sativa variety from B. Seeds Co. and can be cultivated outdoors (where the plants will need a flowering time of ±80 days ) . B. Seeds Co.s AH/PAN/JAM x Sativa is/was never available as feminized seeds.

Amnesia Haze/Panama/Jamaica x Sativa Sudamericana

Flowering time: around 80 days
Untested and limited hybrid, different phenotypes are possible!

Where to buy AH/PAN/JAM x Sativa cannabis seeds?

AH/PAN/JAM x Sativa from B. Seeds Co. is available only as regular seeds. Feminized seeds are not available at the moment. In 1 seedbanks, we found 2 offers between EUR 10.00 for 5 regular seeds and EUR 20.00 for 10 regular seeds. If you are looking to buy AH/PAN/JAM x Sativa Cannabis Seeds from B. Seeds Co. somewhere – have a look to our AH/PAN/JAM x Sativa Price Comparison page with all current offers from all the connected seedbanks and shops – or visit one of the following tested, trustworthy and recommended seed-shops directly to check out their current AH/PAN/JAM x Sativa offers: myCannapot.

AH/PAN/JAM x Sativa Lineage / Genealogy

  • AH/PAN/JAM x Sativa »»» x Unknown Sativa South American
  • Amnesia Haze x Jamaica/Panama
    • Amnesia Haze
      • »»» Afghani Hawaiian x Laos x Jamaica
      • Afghani Hawaiian
        • »»» Afghani x Hawaii
        • Afghani
          • Afghanistan Probably Indica »»» Indica
        • Hawaii »»» Indica/Sativa Hybr >South American »»» Sativa

        Map of the AH/PAN/JAM x Sativa Family Tree

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        Ah/pan/jam x sativa seeds
        Vertical transmission of seed-borne bacterial and fungal endophytes has been detected in various plant species. By isolating Bacillus spp. and Microbacterium spp. from switch grass seeds harvested in 1 year and from the plants grown from these seeds the following year, showed that the same microbial species occurred in multiple (Gagne-Bourgue et al., 2013). In addition, Ringelberg et al. (2012) isolated the same endophytic bacterial genera from both seeds and mature plant tissues in wheatgrass, therefore suggesting that the seeds are a key source of transmitting mature wheatgrass endophytes to the next generation. Furthermore, although fungal endophytes were originally thought to be horizontally transmitted, their vertical transmission in various plant species has been reported (Ngugi and Scherm, 2006; Hodgson et al., 2014; Wiewióra et al., 2015). Some studies indeed report that the rate of vertical transmission for many fungal endophytes is greater than 90% (Ngugi and Scherm, 2006).

        What Is There in Seeds? Vertically Transmitted Endophytic Resources for Sustainable Improvement in Plant Growth

        Raheem Shahzad

        1 School of Applied Biosciences, Kyungpook National University, Daegu, South Korea

        Abdul L. Khan

        2 Chair of Oman’s Medicinal Plants and Marine Natural Products, University of Nizwa, Nizwa, Oman

        Saqib Bilal

        1 School of Applied Biosciences, Kyungpook National University, Daegu, South Korea

        Sajjad Asaf

        2 Chair of Oman’s Medicinal Plants and Marine Natural Products, University of Nizwa, Nizwa, Oman

        In-Jung Lee

        1 School of Applied Biosciences, Kyungpook National University, Daegu, South Korea


        Phytobeneficial microbes, particularly endophytes, such as fungi and bacteria, are concomitant partners of plants throughout its developmental stages, including seed germination, root and stem growth, and fruiting. Endophytic microbes have been identified in plants that grow in a wide array of habitats; however, seed-borne endophytic microbes have not been fully explored yet. Seed-borne endophytes are of great interest because of their vertical transmission; their potential to produce various phytohormones, enzymes, antimicrobial compounds, and other secondary metabolites; and improve plant biomass and yield under biotic and abiotic stresses. This review addresses the current knowledge on endophytes, their ability to produce metabolites, and their influence on plant growth and stress mitigation.


        Soil hosts a diverse array of microbes, such as bacteria, fungi, yeasts, and protozoa. These microbes often exist in mutualistic interactions; some are also found in mutual relationships with plants (Farrer and Suding, 2016; Vejan et al., 2016; Lladó and Baldrian, 2017). These plant–microbe associations have been the focus of comprehensive study, given their potential as ecologically sound alternatives for promoting crop growth and development. It is clear that microorganisms are able to enhance plant growth and defenses, and that plants have the ability to select a microbiome in order to retain valuable colonizers, including those living within their tissue (Hardoim et al., 2012; Marasco et al., 2012; Rashid et al., 2012). Within this context, seed microbiota are ecologically interesting in that they represent not only an endpoint for the community assembly in the seed, but also a starting point for community assembly in the new seedling.

        The present review concentrates on underexplored endophytes, such as seed-borne bacterial and fungal endophytes. The review considers their role in enhancing crop efficiency, the nature of vertical transmission and secondary metabolite production, their below-ground function, and the above-ground response.

        Endophytic microbes: role and reproduction

        Recent evaluations suggest that over 300,000 plant species are found worldwide, and that every plant carries at least one endophyte (Smith et al., 2008). Indeed, endophytic microbes have been found in every plant species examined to date; Partida-Martínez and Heil (2011) report that a plant without endophytes could only occur infrequently. It can be assumed that plants deprived of endophytes would be more vulnerable to environmental stress and pathogenic attacks (Khan et al., 2015; Leitão and Enguita, 2016; Suman et al., 2016; Brader et al., 2017). Endophytic microorganisms (bacteria or fungi) are a key class of plant symbionts that live inside plant tissues without inducing any disease symptoms (Brader et al., 2017), and which are associated with the plant throughout its life history, from seed germination to fruit development. Endophytes are found in the roots (rhizosphere), leaves (phylloplane), stems (laimosphere and caulosphere), fruits (carposphere), seeds (spermosphere), and flowers (anthosphere), as described by many scientists (Clay and Holah, 1999; Lindow and Brandl, 2003; Saikkonen et al., 2004; Shahzad et al., 2016; Brader et al., 2017). The relationship between endophytes and plants is unique in the ability of the former to provide alternative sources of biologically active metabolites, such as enzymes, biofunctional chemicals, phytohormones, nutrients, and minerals, and to facilitate the distribution or production of these resources which contributed in the elimination of various stresses (Schulz et al., 2002; Khan et al., 2012; Kong and Glick, 2017; Nelson, 2017). In return, the host plant provides a protective sanctuary for the microbes within the plant tissues, in which they can grow and reproduce, but without compromising the plant’s own growth resources (Khan et al., 2015).

        Why are endophytes in seeds important?

        Seeds play an important role in the life cycle of spermatophytes; they have the ability to exist in a torpid state for a considerable length of time until growth conditions are suitable, and then develop into a new plant (Nelson, 2004; Geisen et al., 2017). It is probable that seeds benefit from seed-borne bacterial and fungal endophytes, which are thought to promote seed conservation and facilitate seed germination in soil (Chee-Sanford et al., 2006; Rodríguez et al., 2017; Shearin et al., 2017). Seed-borne endophytes are of particular importance because they are passed between successive plant generations via vertical transmission, thus ensuring their presence in the next generation of seedlings (Cope-Selby et al., 2017; Shade et al., 2017). This process of vertical transmission results in a weakening of microbial pathogenic strength in order to support plant growth and development. This mutualism supports and enhances plant survival and microbial proliferation (Rudgers et al., 2009). Moreover, alongside their vital role in plant growth and defense, these seed-borne bacterial and fungal endophytes benefit the host plants through providing their offspring with valuable endosymbionts (Shade et al., 2017).

        Biodiversity trove in seeds

        The internal environment of a seed changes during maturation, which consequently affects the seed endophytic community (Mano et al., 2006). The ability to reside in a seed and adapt to severe environmental conditions are special characteristics of seed endophytes that are rarely found in endophytes isolated from roots, shoots, or other plant tissues. Seed endophytes have the ability to form endospores, thus providing protection from changing conditions inside the seed (Mano et al., 2006; Compant et al., 2011; Kane, 2011). They also maintain other features, such as cell motility and phytase activity, in order to be able to migrate freely inside the plant and enter the seeds before they harden. There have however been relatively few studies examining biodiversity in seed-borne endophytes.

        Seed-borne bacterial endophytes

        Table 1

        Endophytic microbes isolated and characterized from the seeds of different plants.

        Endophytic microbes Function References
        Oryza sativa Paenibacillus polymyxa Glucanase production, anti-phytopathogenic microbe Liu et al., 2017
        Cucumis melo Proteobacteria, Frimicutes, Actinobacteria Glassner et al., 2017
        Oryza sativa Micrococcus yunnanensis, Micrococcus luteus, Enterobacter soli, Leclercia adecarboxylata, Pantoea dispersa, Staphylococcus epidermidis IAA production, plant growth promotion Shahzad et al., 2017c
        Oryza sativa Enterobacter asburiae, Pantoea dispersa, Pseudomonas putida IAA production, phosphate-solubilizing, antifungal, plant growth promotion Verma et al., 2017
        Phragimates australis P. fluorescens, Psedomonas sp., Pantoea sp., Enterobacter sp. Phosphorus-solubilizing, protease production, anti-fungal, plant growth promotion White et al., 2017
        Triticum aestivum Panibacillus sp., Pantoea sp., Bacillus sp. IAA production, antifungal, siderophore production, phosphate-solubilizing, plant growth promotion Díaz Herrera et al., 2016
        Tylosema esculentum Massilia, Kosakonia, Pseudorhodoferax, Caulobacter, Pantoea, Sphingomonas, Burkholderia, Methylobacterium, Bacillus sp., Curtobacterium, Microbacterium, Mucilaginibacter, Chitinophaga Plant growth promotion, phytohormone and metabolite production Chimwamurombe et al., 2016
        Oryza sativa Bacillus amyloliquefaciens Phytohormone production, growth promotion Shahzad et al., 2016
        Lycopersicum esculentum Bacillus subtilis Plant growth promotion, phytohormone and metabolite production Xu et al., 2014
        Zea mays Undibacterium, Sphingomonas, Acinetobacter, Burkholderia, Pantoea, Limnobacter, Burkholderia, Pantoea, Staphylococcus, Serratia, Cronobacter, Enterobacter, Escherichia, Acinetobacter Liu et al., 2013
        Arachis hypogaea B. thuringiensis, B. cereus, B. amyloliquefaciens, B. megaterium, B. subtilis, Bacillus sp., Paenibacillus sp., Pseudomonas sp., B. thioparans, Cyanobacterium Antifungal Sobolev et al., 2013
        Phaseolus vulgaris Bacillus massilensis, Bacillus sp. Bacillus pumilus, Bacillus flexus, Bacillus korlensis, Bacillus silvestris, Paenibacillus, Enterococcus, Staphylococcus, Arthrobacter, Kocuria, Micrococcus, Brachybacterium, Methylobacterium, Paracoccus, Acinetobacter Rosenblueth et al., 2012
        Triticum aestivum; Elymus trachycaulus; Agropyron fragile Actinobacteria, Firmicutes, Gammaproteobacteria Ringelberg et al., 2012
        Oryza sativa Pseudomonas protegens, Pseudomonas sp., Stenotrophomonas maltophilia, Uncultured Stenotrophomonas clone, Ochrobactrum tritici, Ochrobactrum sp., Ochrobactrum grignonense Sphingomonas yanoikuyae, Flavobacterium johnsoniae, Flavobacterium sp., Paenibacillus humicus, Paenibacillus sp. Agromyces mediolanus, Curtobacterium citreum, Curtobacterium sp., Curtobacterium herbarum, Frigoribacterium faeni, Microbacterium oleivorans, Microbacterium sp., Mycobacterium abscessus Plantibacter flavus Plant growth promotion, mitigating biotic and abiotic stress Hardoim et al., 2012
        Zea mays Bacillus sp, Methylobacterium, Tukamurella, Alcaligenes, Erwinia, Microbacterium, Rhodococcus Rosenblueth et al., 2012
        Cucurbita pepo Bacillus sp., Pseudomonas chlororaphis, Lysobacter gummosus, P. chlororaphis, Paenibacillus polymyxa, Serratia plymuthica Antifungal Fürnkranz et al., 2012
        Vitis vinifera Bacillus altitudinis, Bacillus simplex, Bacillus thuringiensis, Paenibacillus amylolyticus, Staphylococcus aureus subsp. aureus Tissue colonization Compant et al., 2011
        Fraxinus Pantoea agglomerans, Staphylococcus succinus, Aerococcus viridans Antibiotic production Donnarumma et al., 2011
        Oryza sativa Pantoea agglomerans, Acinetobacter sp., Curtobacterium citreum, Microbacterium sp., Pantoea ananatis, Pseudomonas sp., Paenibacillus sp., Pantoea sp., Staphylococcus cohnii, Curtobacterium citreum, Microbacterium sp., Sphingomonas sp., Rhizobium larrymoorei, Curtobacterium sp., Sphingomonas sp. Phytohormone and metabolite production, phosphate-solubilizing, antifungal, plant growth promotion Ruiza et al., 2011
        Glycine max Acinetobacter, Bacillus, Enterococcus, Nocardioides, Paracoccus, Phyllobacterium, Sphingomonas Phytate-solubilizing López-López et al., 2010
        Nicotiana tabacum Enterobacter sp., Xanthomonadaceae, Pseudomonas sp., Enterobacter sp., Pseudomonas fulva, Sanguibacter sp., Stenotrophomonas sp., Clostridium aminovalericum, Stenotrophomonas sp., Sanguibacter sp. Mitigating metal toxicity, promote plant growth Mastretta et al., 2009
        Oryza sativa Bacillus pumilus, Kocuria palustris, Pantoea ananatis, Methylobacterium radiotolerans, Methylobacterium fujisawaense Enzyme production, osmotic stress tolerance Kaga et al., 2009
        Eucalyptus Bacillus sp., Enterococcus sp., Paenibacillus sp., Methylobacterium sp. Growth promotion Ferreira et al., 2008
        Zea mays Pantoea sp., Microbacterium sp., Frigoribacterium sp., Bacillus sp., Paenibacillus sp., Sphingomonas sp. Antifungal Rijavec et al., 2007
        Oryza sativa Xanthomonas translucens, Pantoea ananatis, Methylobacterium aquaticum, Sphingomonas melonis, Sphingomonas yabuuchiae, Bacillus subtilis, Bacillus pumilus, Micrococcus luteus, Acidovorax sp., Curtobacterium flaccumfaciens, Paenibacillus amylolyticus, Xanthomonas translucens Enzyme production, osmotic stress tolerance Mano et al., 2006
        Coffea Arabica Bacillus sp., Burkholderia cepacia—GC subgroup B, Burkholderia gladioli GC subgroup A, Burkholderia gladioli—GC subgroup B, Clavibacter michiganense insidiosum, Curtobacterium flaccumfaciens-flaccumfaciens, Curtobacterium flaccumfaciens-poinsettiae, Escherichia vulneris, Micrococcus sp., Pantoea agglomerans, Pseudomonas putida biotype A, Pseudomonas putida biotype B, Stenotrophomonas sp., Stenotrophomonas maltophilia, Yersinia frederiksenii Vega et al., 2005
        Fragaria Pseudomonas fluorescens, Pseudomonas sp. Kukkurainen et al., 2005
        Glycine max Agrobacterium radiobacter, Aeromonas sp., Bacillus spp., Chryseomonas luteola, Flavimonas oryzihabitans, Sphingomonas paucimobilis Seedling growth, root colonization Oehrle et al., 2000
        Invasive Phragmites Alternaria sp., Phoma sp., Penicillium corylophilum Improved seed germination and seedling growth Shearin et al., 2017
        Dendrobium friedericksianum Fusarium sp., Beauveria sp., Tulasnella violea, T. violea, Epulorhiza sp., Trichosporiella multisporum Growth promotion Khamchatra et al., 2016
        Cinchona ledgeriana Diaporthe sp. Alkaloid production Maehara et al., 2016
        Toona sinensis Roem Cladosporium sp. Antioxidant potential Rahmawati et al., 2016
        Lolium perenne Neotyphodium sp. Wiewióra et al., 2015
        Schedonorus phoenix Epicholë ceonophiala Improved resistance against herbivores and environmental stresses Young et al., 2013
        Dactylis glomerata Epichloë typhina Improved host plant growth and photosynthesis Rozpadek et al., 2015
        Centaurea cyanus;
        Papaver rhoeas;
        Senecio vulgaris;
        Centaurea nigra;
        Plantago lanceolata;
        Rumex acestosa
        Acremonium strictum, Alternaria alternate, Aspergillus niger, Aureobasidium pullulans, Botrytis cinerea, Chaetomium cochliodes, Clodosporium cladospriodes, Cladosporium oxysporum, Cladosporium sphaerospermum, Colletotrichum dematium, Epicoccum nigrum, Fusarium avenaceum, Fusarium equiseti, Fusarium merismoides, Fusarium tricinctum, Fusarium sp. A, Geotrichum candidum, Mucor hiemalis, Penicillium sp A, Penicillium sp. B, Phialophora verrucosa, Rhabdospora coricea, Sterile sp. A, Sterile sp. B Hodgson et al., 2014
        Laelia speciosa Helotiales sp. Ávila-Díaz et al., 2013
        Ipomoea carnea Collelotrichum sp., Fusarium sp. Antimicrobial Tayung et al., 2012
        Swietenia macrophylla King α –Glucosidase inhibition Ramdanis et al., 2012
        Festuca arundinacea Neotyphodium oenophialum Ergovaline and loline alkaloid production and improved protection against herbivores Pennell et al., 2010
        Lolium perenne Epichloë festucae var. lolii Improved drought tolerance Kane, 2011

        Seed-borne fungal endophytes

        Fungal endophytes are found in all types of plant tissue, and have been shown to improve growth, enhance plant defense systems, and mitigate both biotic and abiotic stress (Khan et al., 2015). Endophytic fungi reveal a broad variation in their mode of transmission from one host to another, and stringent vertical transmission from one generation to the next (Shearin et al., 2017; Vujanovic and Germida, 2017). Many fungi are seed-borne, and very recent studies report that fungal seed microbiomes may be greatly influenced by local conditions and non-host genotypes (Klaedtke et al., 2016). The well-studied seed-borne fungal endophytes belonging to the genus Epichlöe are mostly reported to assist their host plants in growth promotion and stress mitigation, either directly or indirectly (Kauppinen et al., 2016; Gundel et al., 2017). However, although research has focused on this group of fungi, there are numerous other seed-associated fungi, including ascomycetes, basidiomycetes, parasites, and yeasts (Abe et al., 2015).

        Mechanisms of action of seed-borne endophytes

        The assorted metabolic qualities of seed-borne bacterial and fungal endophytes are dependent on local conditions, and are used to facilitate the host plant’s advancement. This further strengthens the benefits conferred on the host plant, improving its fitness over other plants; this can in turn influence the entire environment (Klironomos, 2002; Khan et al., 2015; Figure ​ Figure1A 1A ).

        Conceptual view of mechanisms of action and vertical transmission of seed endophytic microbiota. (A) The schematic presentation shows the isolation of seed-borne endophytic microbes, and their application in promoting plant growth and stress resistance. (B) A holistic view of the vertical transmission of seed-borne endophytes. This suggests that endophytes are found in seed embryos and grow into the emerging leaf upon germination; the endophytes then migrate into the stem and seed head of reproductive plants via various pathways.

        Mode for vertical transmission of seed-borne endophytes

        Vertical transmission of seed-borne bacterial and fungal endophytes has been detected in various plant species. By isolating Bacillus spp. and Microbacterium spp. from switch grass seeds harvested in 1 year and from the plants grown from these seeds the following year, showed that the same microbial species occurred in multiple (Gagne-Bourgue et al., 2013). In addition, Ringelberg et al. (2012) isolated the same endophytic bacterial genera from both seeds and mature plant tissues in wheatgrass, therefore suggesting that the seeds are a key source of transmitting mature wheatgrass endophytes to the next generation. Furthermore, although fungal endophytes were originally thought to be horizontally transmitted, their vertical transmission in various plant species has been reported (Ngugi and Scherm, 2006; Hodgson et al., 2014; Wiewióra et al., 2015). Some studies indeed report that the rate of vertical transmission for many fungal endophytes is greater than 90% (Ngugi and Scherm, 2006).

        Metagenome analysis

        Seeds are not merely the carriers of a plant’s hereditary information, but also both reservoirs for plant microbiota and vehicles for their vertical transmission (Baker and Smith, 1966; Nelson, 2004). The role of seed-associated microbes is of significance to plant growth and development because these microbial communities may secrete important phytohormones, such as cytokinins, that break seed dormancy (Goggin et al., 2015) and inhibit microbial invasions (Bacilio-Jiménez et al., 2001). The recent rapid progress in high-throughput DNA sequencing technology has enabled a far wider exploration of microbes in the rhizosphere, endosphere, and phyllosphere of important crops and model plant species, revealing the distinctive microbial community structures, which are dependent on the plant parts they inhabit and environmental conditions (Redford and Fierer, 2009; Bulgarelli et al., 2012; Bodenhausen et al., 2013; Shakya et al., 2013; Lebeis, 2014). Furthermore, multi-omics techniques, such as whole genome and metagenomic analyses, have significantly improved our understanding of the role of the plant microbiome (Bai et al., 2015; Bulgarelli et al., 2015).

        Extensive attention has been given to the construction and role of microbial communities associated with the phyllosphere and rhizosphere. However, we have a comparatively poor understanding of the microbiota inhabiting other niches, such as the reproductive organs and seeds. Seeds form an important habitat for microbes, sustaining a diverse array of both harmful and beneficial microbes (Nelson, 2004). Similar to the rhizosphere, the spermosphere is a region that surrounds seeds, and in which seed microbes, germinating seeds, and soil microbes may interact (Nelson, 2004). The microbiota living in this region, although usually short-lived as individual organisms, can have a persistent effect on seed germination and seedlings (Nelson, 2004; Delgado-Sánchez et al., 2011; Chen et al., 2012; Schiltz et al., 2015). Recently, research has revealed that microbes in the seed spermosphere and endosphere, which are less studied than other groups of symbionts, have the ability to promote seed germination and enhance plant growth during both abiotic and biotic stress (Truyens et al., 2015). For example, fungi isolated from Opuntia spp. (Penicillium chrysogenum, Phoma sp., and Trichoderma koningii) are involved in breaking seed dormancy and promoting germination (Delgado-Sánchez et al., 2011, 2013). Similarly, some seed-borne endophytic fungi from Ascomycota and Pleosporales have been reported to promote the growth and germination of Phragmites australis (Ernst et al., 2003). In addition, the effects of seed-associated microbiota on seed germination and plant growth are not limited to plant–fungal interactions; seed-associated bacteria have also been found to have similar functions in relation to plant fitness (Xu et al., 2014; Hardoim et al., 2015; Pitzschke, 2016). Therefore, it is reasonable to hypothesize that seed-associated microbes, including epiphytes and endophytes, play a more important role in modulating their host plant than previously thought.

        High-throughput sequencing studies have identified a high frequency of Cladosporium spp. in seeds, specifically the inner seeds of a wide range of herbaceous plants (Ikeda et al., 2006; Lucero et al., 2011). Similarly, it has been reported that both endophytes and epiphytes associated with seeds play significant roles in seed germination and plant growth (Pitzschke, 2016; Tahtamouni et al., 2016). In rainy tropics, seed epiphytic fungi (Penicillium sp. and Fusarium sp.) have been shown to enhance seed germination (Tamura et al., 2008). Thus, exploration of these microbial communities using modern metagenomics has revealed there to be genetic and biochemical diversity in the spermosphere and endosphere of seeds.

        Plant growth promotion and stress tolerance

        Although there has been a w >​ (Table1). 1 ). Several seed-borne bacterial and fungal endophytes produce compounds that either directly inhibit pathogen growth or indirectly strengthen plant resistance in defense against pathogenic attack (Bonos et al., 2005; Clarke et al., 2006; Tayung et al., 2012; Shahzad et al., 2017a). Yue et al. (2000) have determined the occurrence of numerous indole compounds, a sesquiterpene, and diacetamide from Epichloë festucae. Moreover, Shahzad et al. (2017a) report that the various organic acids produced by seed-borne endophytic Bacillus amyloliquefaciens acted to significantly inhibit the growth of pathogenic Fusarium oxysporum in vitro, and induced systemic resistance in tomato plants. Díaz Herrera et al. (2016) report the isolation from wheat seeds of the endophytes Paenibacillus sp., Pantoea sp., and Bacillus sp., which significantly enhanced plant growth and resistance against F. graminearum. Furthermore, Epicholë grass endophytes are also widely used in improving the survival and productivity of perennial ryegrass (Karpyn Esqueda et al., 2017). Turfgrasses infected with E. festucae showed a significantly improved resistance in comparison with non-inoculated turfgrasses against two of the main leaf spot pathogens, Sclerotina homeocarpa and Laetisaria fuciformis (Bonos et al., 2005; Clarke et al., 2006). However, it remains unclear whether this enhanced defense mechanism is attributable to metabolites produced by endophytes, secondary metabolites produced by plants in response to inoculation by endophytes, or competition between pathogenic microbes. Interestingly, in addition to their antagonistic capability against pathogenic microbes, seed endophytes also improve seed germination, mitigate abiotic stress, and enhance plant tolerance, features which are probably related to the ability of these microbes to produce secondary metabolites, siderophores, and ACC deaminase (Glick, 2012; Xu et al., 2014; Shahzad et al., 2017a,b). Moreover, the application of plant growth-promoting seed-borne bacterial endophytes may also facilitate the phyto- and bioremediation of contaminated soil. Mastretta et al. (2009) have shown in their study that, inoculation of tobacco plants with seed endophytes under Cd stress resulted in significantly improved plant growth, enhanced biomass, alleviation of Cd toxicity, and improved tolerance as compared to uninoculated plants. Truyens et al. (2015) also report enhanced phytoremediation of grasses following inoculation with seed-borne endophytes with the potential to solubilize phosphorus and produce indole-3-acetic acid (IAA), siderophores, ACC deaminase, and acetone. They also conclude that there are benefits to establishing Cd-tolerant seed-borne endophytes in Cd-contaminated areas during phytoextraction and phytostabilization; in non-exposed plants, endophyte inoculation considerably improved plant growth, whereas under conditions of Cd stress, inoculation augmented Cd uptake without disturbing plant growth. These results show that endophyte microbes such as these are promising in terms of applicability to phytoremediation.

        Future perspectives

        Investigating the role of seed-borne, vertically transmitted bacterial and fungal endophytes opens new and exciting opportunities for applied research into plant–microbe interactions, given that these microbes can improve seed germination, promote seedling health, enhance plant growth, and mitigate stress. These abilities can be attributed to the production of extracellular enzymes, phytohormones, and secondary metabolites. Given the growth-promoting and biocontrol properties of these microbes, their potential applications as biofertilizers and in bioremediation should be supported.

        It is presumed that an extensive proportion of the endophytic population in seeds has not yet been fully explored. Metagenomic studies will provide additional insight into seed endophyte populations, including the genera, their phenotypic attributes, and possible roles in both germination and plant advancement. Further research is required in order to investigate seed–endophyte interactions and their role in inducing defense resistance mechanisms against biotic at the molecular level and also to identify the genetic determinants involved in seed colonization, seed endophyte dispersal, and vertical transmission. Finally, exhaustive research is needed to determine the changes that occur in seed-associated endophytes during seed development, storage, and germination, in order to ensure a superior quality production of seeds.

        Author contributions

        RS design the study. RS, AK, SB, and SA wrote the review manuscript. I-JL and AK critically reviewed the manuscript and supervised the manuscript drafting.

        Conflict of interest statement

        The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.


        This study was supported by the Basic Science Research Program through the National Research Foundation of Korea (NRF), funded by the Ministry of Education (2017R1D1A1B04035601).