Cabay RJ, Morton TH Jr, Epstein JB. Published by Elsevier Inc. All rights reserved. 2016 Sep;43(9):759-65. doi: 10.1111/cup.12738. Mignogna MD, Fortuna G, Leuci S, et al. Angulated tumor nests infiltrate the underlying stroma. Epub 2020 Oct 16. 2007;36(5):25561. TP53; atypical squamous nodule; keratoacanthoma; p53; squamous cell carcinoma; squamous proliferation. 1985;60(3):28598. Although VV (common wart) usually occurs on extragenital skin, it can occasionally arise in the anogenital area and may result in diagnostic confusion with condyloma acuminatum. [54], Plants are potentially able to break down or sequester, or stimulate micro-organisms in the soil to break down or sequester, certain organic pollutants and inorganic pollutants such as nickel in degraded ecosystems, which is known as phytoremediation. Cases associated with high-grade intraepithelial neoplasia and warty/basaloid squamous cell carcinoma (SCC) are coinfected with high-risk HPV types.68. When verrucous carcinoma forms on your feet, they manage it as a skin cancer. which was first harvested from the plant Bontia daphnoides. The eruption was initially treated with topical 5-FU twice daily for 4 weeks to treat his large hyperkeratotic lesions on the lower legs that were concerning for multifocal eruptive squamous atypia Contrary to historical belief, giant condyloma acuminatum and verrucous carcinoma should be considered distinct entities based on different pathogenetic pathways. Med Oral Patol Oral Cir Bucal. Clavicipitaceous and non-clavicipitaceous, "An Archaic Approach to a Modern Issue: Endophytic Archaea for Sustainable Agriculture", "The Hidden World within Plants: Ecological and Evolutionary Considerations for Defining Functioning of Microbial Endophytes", "Defining the Genetic Basis of PlantEndophytic Bacteria Interactions", Creative Commons Attribution 4.0 International License, "Exploring the evolutionary ecology of fungal endophytes in agricultural systems: using functional traits to reveal mechanisms in community processes", "Experimental evidence of microbial inheritance in plants and transmission routes from seed to phyllosphere and root", "A Friendly Relationship between Endophytic Fungi and Medicinal Plants: A Systematic Review", "Fungal endophytes and their interaction with plant pathogens: a review", "In planta regulation of extension of an endophytic fungus and maintenance of high metabolic rates in its mycelium in the absence of apical extension", "Chemical ecology of endophytic fungi: origins of secondary metabolites", "Increased Biomass of Nursery-Grown Douglas-Fir Seedlings upon Inoculation with Diazotrophic Endophytic Consortia", "An In vitro Study of Bio-Control and Plant Growth Promotion Potential of Salicaceae Endophytes", "Endophyte Effects on Photosynthesis and Water Use of Plant Hosts: A Meta-Analysis", "Salicaceae Endophytes Modulate Stomatal Behavior and Increase Water Use Efficiency in Rice", "Root ethylene signalling is involved in Miscanthus sinensis growth promotion by the bacterial endophyte Herbaspirillum frisingense GSF30T", "Endophytic root colonization of gramineous plants by Herbaspirillum frisingense", "Fungal endophytes: diversity and functional roles", "Mycorrhizal ecology and evolution: the past, the present, and the future", "Bacterial endophytes in agricultural crops and their role in stress tolerance: a review", "Bioprospecting for microbial endophytes and their natural products", "Endophytic fungi: a reservoir of antibacterials", "Can naturally-occurring endophytic nitrogen-fixing bacteria of hybrid white spruce sustain boreal forest tree growth on extremely nutrient-poor soils? [20] Specific endophyte species can also stimulate root growth by increasing the flux of auxin to where the endophyte is. PubMed Central [28] One group of fungal endophytes are the arbuscular mycorrhizal fungi involving biotrophic Glomeromycota associated with various plant species. The endophytes can survive most pesticides and are even resistant to some fungicides, and are very suitable for use in Integrated Pest Management. Architectural Alterations in Oral Epithelial Reverse maturation and stromal invasion, when present, should raise concern for malignant transformation.21,27 Immunohistochemical profile, including negative, focal, or patchy staining for p16, is the same as described in ordinary condyloma acuminatum.24, Verrucous carcinoma is a unique variant of well-differentiated SCC. There are two different means of classifying endophytes. Arch Pathol Lab Med 1 July 2019; 143 (7): 821831. An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Oral Surg Oral Med Oral Pathol Oral Radiol. The adjacent epidermis is acanthotic but not dysplastic. Manchanda A, Shetty DC. Head Neck Pathol. Am J Dermatopathol. The working group then selected 43 single slide biopsy cases for whole slide digital imaging (WSI) review by members of the consensus conference. These interaction range from pathogenic to symbiotic and symptomatic to asymptomatic. In another experiment Burkholderia bacteria with both the TOM plasmid as well as nickel resistance genes was inoculated into yellow lupine; this increased the root mass of the plants, but the amounts of TCE transpired was not statistically significant. Skin | SpringerLink HPV infection and immunochemical detection of cell-cycle markers in verrucous carcinoma of the penis, The association of lichen planus of the penis with squamous cell carcinoma in situ and with verrucous squamous carcinoma, Prevalence of mucosal and cutaneous human papillomaviruses in different histologic subtypes of vulvar carcinoma, Verrucous carcinoma of the skin and mucosa, Surgical treatment for 11 cases of penile verrucous carcinoma, Verrucous carcinoma: clinical and pathologic study of 105 cases involving oral cavity, larynx and genitalia, Diagnostic problems in precancerous lesions and invasive carcinomas of the penis, Primary squamous cell carcinoma of the vagina: human papillomavirus detection, p16(INK4A) overexpression and clinicopathological correlations, Worldwide human papillomavirus genotype attribution in over 2000 cases of intraepithelial and invasive lesions of the vulva, Prevalence and distribution of 15 high-risk human papillomavirus types in squamous cell carcinoma of the scrotum, Histologic classification of penile intraepithelial neoplasia, The 2015 International Society for the Study of Vulvovaginal Disease (ISSVD) terminology of vulvar squamous intraepithelial lesions, The Lower Anogenital Squamous Terminology standardization project for HPV-associated lesions: background and consensus recommendations from the College of American Pathologists and the American Society for Colposcopy and Cervical Pathology, Distribution and characterization of subtypes of penile intraepithelial neoplasia and their association with invasive carcinomas: a pathological study of 139 lesions in 121 patients, Warty (condylomatous) squamous cell carcinoma of the penis: a report of 11 cases and proposed classification of verruciform' penile tumors, Squamous cell carcinoma of the vulva: a subclassification of 97 cases by clinicopathologic, immunohistochemical, and molecular features (p16, p53, and EGFR), p16 overexpression identifies HPV-positive vulvar squamous cell carcinomas, Distinctive association of p16INK4a overexpression with penile intraepithelial neoplasia depicting warty and/or basaloid features: a study of 141 cases evaluating a new nomenclature, Distinctive immunohistochemical profile of penile intraepithelial lesions: a study of 74 cases, Anal intraepithelial neoplasia: a single centre 19 year review, Papillary squamous cell carcinoma, not otherwise specified (NOS) of the penis: clinicopathologic features, differential diagnosis, and outcome of 35 cases, Bowenoid papulosis: a clinicopathologic study with ultrastructural observations, Bowenoid papulosis: clinical and histological study of eight cases, Bowenoid papulosis: presence of human papillomavirus (HPV) structural antigens and of HPV 16-related DNA sequences, Bowenoid papulosis of the vulva-immunotherapeutical approach with topical imiquimod, Photodynamic therapy for bowenoid papulosis using a novel incoherent light-emitting diode device, Extensive genital bowenoid papulosis responding to imiquimod, Aminolevulinic acid photodynamic therapy for bowenoid papulosis, Bowenoid papulosis: classification as a low-grade in situ carcinoma of the epidermis on the basis of histomorphologic and DNA ploidy studies, Bowenoid papulosis of the male and female genitalia: risk of cervical neoplasia, Heterogeneity of human papillomavirus DNA in a patient with Bowenoid papulosis that progressed to squamous cell carcinoma, Morphometric assessment of nuclei in Bowen's disease and bowenoid papulosis, The presence of HPV types 6/11, 16/18, 31/33/51 in Bowenoid papulosis demonstrated by DNA in situ hybridization, Detection of mucosal human papilloma virus DNA in bowenoid papulosis, Bowen's disease and squamous cell carcinoma of the skin, Expression of p16 and hTERT protein is associated with the presence of high-risk human papillomavirus in Bowenoid papulosis, A genotype distribution of human papillomaviruses detected by polymerase chain reaction and direct sequencing analysis in a large sample of common warts in Japan, Development of a multiplex PCR method for detecting and typing human papillomaviruses in verrucae vulgaris, Verruca vulgaris of the vulva in children and adults: a nonvenereal type of vulvar wart, Human papillomavirus segregation patterns in genital and nongenital warts in prepubertal children and adults, Immunohistochemistry and polymerase chain reaction for detection human papilloma virus in warts: a comparative study, Multiple epidermolytic acanthomas must not be confused with genital human papillomavirus infection, Epidermolytic acanthoma of the scrotum: a rare mimicker of condyloma acuminatum, Multiple localized epidermolytic acanthomas of the vulva associated with vulvar pruritus: a case report, Multiple epidermolytic acanthomas mimicking condyloma: a retrospective study of 8 cases, Epidermolytic acanthoma does not contain human papillomavirus DNA. Oral carcinoma cuniculatum: an unacquainted entity with and have shown cytotoxicity against human leukemia (K-562). Biopsy of a lesion in the right proximal pretibial region demonstrated an atypical endophytic squamous proliferation, consistent with SCC. [68] Epichlo endophytes are being widely used commercially in turf grasses to enhance the performance of the turf and its resistance to biotic and abiotic stresses. [55], Two strains of the endophytic fungi Pestalotiopsis microspora isolated from stems of plants from the Ecuadorian rainforest were shown in laboratory experiments to be able to digest polyurethane plastic as the fungus's sole carbon source in anaerobic conditions, although many other non-endophytic fungi have demonstrated this ability, and most isolates of endophytic fungi in this experiment could perform this to some degree. Verruciform and Condyloma-like Squamous C, An anogenital seborrheic keratosis with an undulating base and numerous pseudohorn cysts without evidence of koilocytosis. Histologic distinction between condyloma acuminatum and various benign and malignant condyloma-like lesions in the anogenital area poses a common diagnostic challenge to pathologists across subspecialties. 2017;83(5):58593. This is a bulky squamous proliferation, three to four times the thickness of the epithelium for the site, that is endophytic and/or exophytic in its growth pattern In a series of 108 cases, it was concluded that a background of maturing keratinocytes and sparing of the follicular epithelium could distinguish bowenoid papulosis from basaloid HSIL.53 Another morphometric analysis showed significantly larger and more irregular nuclei in basaloid HSIL compared to bowenoid papulosis.64 In practice, however, the histopathologic differences are often subtle, and definitive diagnosis relies heavily on stringent clinical correlation: Presence of multiple small pigmented papules in a young adult, as opposed to a solitary, slowly progressive plaque or erythematous patch in an older patient, would support a diagnosis of bowenoid papulosis over basaloid HSIL.53,60, High-risk HPV in bowenoid papulosis can be detected by in situ hybridization or PCR-DNA sequencing.6567 Expression of both p16 and human telomerase reverse transcriptase (hTERT) by immunohistochemistry is associated with high-risk HPV infection and has been demonstrated in bowenoid papulosis.67 Diffuse p16 expression is especially helpful in differentiating occasional cases of bowenoid papulosis with more orderly maturation, subtle cytologic atypia, and koilocyte-like cells from condyloma acuminatum (Figure 6, B).18. Systemic endophytes are defined as organisms that live within plant tissues for the entirety of its life cycle and participate in a symbiotic relationship without causing disease or harm to the plant at any point. Additionally, systemic endophytes concentrations and diversity do not change in a host with changing environmental conditions. eCollection 2019 Dec. Histopathologic features of multiple cutaneous squamous cell carcinomas of the lower extremity. 2019;96:12130. By continuing to use our website, you are agreeing to, SPECIAL SECTIONESSENTIAL DERMATOPATHOLOGY FOR THE GENERAL SURGICAL PATHOLOGIST, PART I, GIANT CONDYLOMA ACUMINATUM (BUSCHKE-LWENSTEIN TUMOR), WARTY/WARTY-BASALOID HIGH-GRADE SQUAMOUS INTRAEPITHELIAL LESION AND SQUAMOUS CELL CARCINOMA, https://doi.org/10.5858/arpa.2018-0039-RA, Molecular Profiling of Syringocystadenocarcinoma Papilliferum Reveals, Transcription Factor Immunohistochemistry in the Classification of Pituitary Neuroendocrine Tumor/Adenoma, Using Pathology Synoptic Reporting Data to Create Individual Dashboards for Pathologists and Surgeons, TRPS1 and GATA3 Expression in Invasive Breast Carcinoma With Apocrine Differentiation, Navigating Practice Issues Related to the Unsatisfactory Cervicovaginal Papanicolaou Test: Survey Results of Laboratories Participating in the 2020 College of American Pathologists PAP Education Program, Updated Salivary Gland Immunohistochemistry: A Review, The Application and Pitfalls of Immunohistochemical Markers in Challenging Diagnosis of Genitourinary Pathology, The Prevalence of SARS-CoV-2 in Autopsies Surrounding the Time of Pandemic Onset: A Retrospective Review of Cases, MicroRed Blood Cell, Fragmented Red Blood Cell, Platelet Distribution Width, Mean Platelet Volume, and PlateletLarge Cell Ratio on Sysmex XN Series Hematology Analyzers Can Be Used for the Reflex Test of Impedance Platelet Count in Clinical Practice. Montague LJ, Bhattacharyya I, Islam MN, Cohen DM, Fitzpatrick SG. PubMed [56], Endophytes produce a wide variety of secondary metabolites that might be useful as lead compounds in drug discovery. This fungus yields a metabolite that shows potential as an antidiabetic, also known as an insulin mimetic. Similar lesions frequently harbor human papillomavirus and should at least raise suspicion for a seborrheic keratosis-like condyloma acuminatum. A lymphocytic band is present at the tumor base. The latter refers to well-differentiated SCC with a papillary architecture. A review of the literature on condyloma acuminatum (ordinary and giant types), verrucous carcinoma, warty/warty-basaloid high-grade squamous intraepithelial lesion and squamous cell carcinoma, papillary squamous cell carcinoma, bowenoid papulosis, verruca vulgaris, epidermolytic acanthoma, and verruciform xanthoma was performed. Oral Surg Oral Med Oral Pathol Oral Radiol. Plant-microbe interactions are not strictly mutualistic, as endophytic fungi can potentially become pathogens or saprotrophs, usually when the plant is stressed. The elongated rete ridges tend to converge toward the center of the lesion (in-toeing) (Figure 7, A). Akrish S, Eskander-Hashoul L, Rachmiel A, Ben-Izhak O. Clinicopathologic analysis of verrucous hyperplasia, verrucous carcinoma and squamous cell carcinoma as part of the clinicopathologic spectrum of oral proliferative verrucous leukoplakia: a literature review and analysis. Bagan JV, Murillo J, Poveda R, et al. Verruciform xanthomas are well-demarcated, yellowish-orange papules or plaques with verruciform surfaces ranging from 0.2 to 2 cm in diameter.89 Not surprisingly, the clinical morphology often raises concern for condyloma acuminatum or verrucous carcinoma.89,90 In a study of 10 vulvar verruciform xanthomas, all were associated with vulvar inflammatory or neoplastic conditions including lichen sclerosus, lichen planus, Paget disease, and radiodermatitis.89 Although HPV has been proposed in the past as the etiologic agent, multiple studies have failed to confirm a causal relationship.9193 Based on its benign nature and association with various diseases, it is now widely believed that verruciform xanthoma represents a mucocutaneous reaction to localized trauma and chronic inflammation.94. Benign alveolar ridge keratosis (oral lichen simplex chronicus): a distinct clinicopathologic entity. 2016;122(3):33254. Speight PM, Abram TJ, Floriano PN, et al. To facilitate unified nomenclature across subspecialties, the Lower Anogenital Squamous Terminology (LAST) work group has recommended a 2-tier system for all HPV-associated noninvasive squamous lesions of the lower anogenital tract: low-grade squamous intraepithelial lesion (LSIL) and high-grade squamous intraepithelial lesion (HSIL).43 It is optional to further classify LSIL and HSIL as -IN 1 and -IN 2 or 3, respectively, according to the specific sites.43 Although HSIL is synonymous with SCC in situ and Bowen disease, the latter terminologies are best avoided in pathology reporting for the purpose of consistency. Upadhyaya JD, Fitzpatrick SG, Islam MN, Bhattacharyya I, Cohen DM. [32], A diazotrophic bacterium isolated in lodgepole pines (Pinus contorta) in British Columbia, Canada, is Paenibacillus polymyxa, which may help its host by fixing nitrogen.[33][34][35][36][37][38]. J Oral Maxillofac Surg. World Workshop on Oral Medicine VII: Prognostic biomarkers in oral leukoplakia and proliferative verrucous leukoplakiaA systematic review of retrospective studies. [5], Most endophyte-plant relationships are still not well understood. All procedures performed in this retrospective data analysis involving human participants were in accordance with the ethical standards of the institutional review board, which did not require informed consent. Squamous cell papilloma (SCP) is defined as a benign proliferation of the surface epithelium of various organs including the skin, lip, tongue, oral cavity, larynx, [13][14] Some fungal and bacterial endophytes have proven to increase plant growth and improve overall plant hardiness. The lesions demonstrated the spectrum of histologic features described in PVL, and in some cases, patients developed oral cavity squamous cell carcinoma (SCC). Head Neck Pathol. There is no evidence of koilocytosis (hematoxylin-eosin, original magnifications 20 [A] and 200 [B]). A, A small verrucous papule with marked hyperkeratosis. From the Department of Pathology, University of Michigan Health System, Ann Arbor. Oral Oncol. These findings suggest that some cutaneous squamous proliferations on the legs of women with multiple lesions lack prominent cytologic atypia as well as TP53 mutations and might be more akin to keratoacanthoma than SCC or might represent a reactive phenomenon. Bookshelf 2014;117(4):51120. The proliferation barely protrudes above the surface Full size image Hyperkeratotic or papillomatous variant is characterized by prominence of papillomatosis, hyperkeratosis, and horn pseudocysts with some acanthosis (Fig. WebAn endophytic fungus, Pseudocercosporella trichachnicola, was found to be widespread in the warm-season grass species Trichachne insularis ( White et al., 1990 ). As of 1999 this is only available in the afore-mentioned lawn grasses, which are sold as 'low maintenance' cultivars. Fischer DJ, Epstein JB, Morton TH, Schwartz SM. The best inter-observer diagnostic agreement relative to PVL lesions were classified as corrugated ortho(para)hyperkeratotic lesion, not reactive and SCC (chi-square p=0.015). These features should prompt a closer inspection for xanthomatous (foam) cells, which are confined to the dermal papillae (Figure 9, C). Class 4 endophytes are restricted to plant tissues below ground and can colonize much more of the plant tissue. 2014;36(11):16628. 2020. https://doi.org/10.1002/jper.19-0621. PubMed [32] It has been estimated that there could be approximately 1 million endophytic fungi that exist in the world. Besides condyloma acuminatum, a number of cutaneous and mucosal lesions in the anogenital region share a similar verruciform architecture and/or clinical presentation, often representing a common source of diagnostic confusion. [50] Turfgrass seed of Festuca and Lolium perenne infected with fungal inoculants, Acremonium coenophialum and A. lolii, is commercially available for use in growing lawns which might require less pesticide use -the grasses are poisonous to cattle and more resistant to some insect damage. Histopathologic examination confirms an exophytic papillomatous lesion. [57][58] Endophyte bioprospecting has already yielded compounds with antibacterial,[59][60] antifungal,[61] antiviral,[61] antiparasitic,[62] cytotoxic,[59][63] neuroprotective,[62] antioxidant,[62] insulin-mimetic,[62] -glucosidase inhibitory,[61] and immunosuppressant[62] properties. J Cutan Pathol. There was moderate agreement (>0.41 kappa) for 35 of 48 cases. Woo SB, Lin D. Morsicatio mucosae orisa chronic oral frictional keratosis, not a leukoplakia. The case for antiherbivore defences", "Exploitation of endophytes for sustainable agricultural intensification", https://en.wikipedia.org/w/index.php?title=Endophyte&oldid=1141734599, Creative Commons Attribution-ShareAlike License 3.0, This page was last edited on 26 February 2023, at 14:41. Class 1 endophytes typically confer benefits on their plant host such as improving plant biomass, increasing drought tolerance and increasing the production of chemicals that are toxic and unappetizing to animals, thereby decreasing herbivory. Fivenon-PVL histologic mimicswere included as controls. Occasional koilocytes may be found in the upper layers.37 Mitoses and apoptoses are frequent and found at all levels of the epithelium (Figure 4, C; inset).46 When invasion occurs in the basaloid and warty-basaloid subtypes, the invasive nests often take on a basaloid appearance without keratinization. The https:// ensures that you are connecting to the Connolly M, Narayan S, Oxley J, de Berker DA. Vulvar seborrheic keratosis: is there a relationship to human papillomavirus? Objective: 2009;45(45):31723. Its giant variant (giant condyloma acuminatum of Buschke-Lwenstein) is discussed separately in the next section. 2010;15(6):e83945. As humans become more aware of the damage that synthetic insecticides cause to the environment and beneficial insects such as bees and butterflies biological insecticides may become more important to the agricultural industry.[31]. Head Neck. 2006;8(2):6770. 2020;42(3):53955. Accessibility There was a belief that plants were healthy under sterile conditions and it was not until 1887 that Victor Galippe discovered bacteria normally occurring inside plant tissues. Recipient(s) will receive an email with a link to 'Verruciform and Condyloma-like Squamous Proliferations in the Anogenital Region' and will not need an account to access the content. This article reviews the clinicopathologic features and etiopathogenesis of condyloma acuminatum and its mimics, with an emphasis on their morphologic similarities and unique characteristics that help in their distinction. Asian Pac J Cancer Prev. 2007;138(5):64151. Koilocytosis may be subtle but often can be appreciated in a background of hypergranulosis. Characterized by complex, confluent, back to back glandular proliferation, while oncocytic papilloma has a simple exophytic or endophytic growth pattern without architectural complexity High grade tumors may show marked nuclear atypia Nonkeratinizing squamous cell carcinoma: Fibrovascular cores are thin and inconspicuous relative to the hypertrophic epithelium. [69] Piriformospora indica is an interesting endophytic fungus of the order Sebacinales, the fungus is capable of colonising roots and forming symbiotic relationship with many plants. Keratoacanthoma, committed stem cells and neoplastic aberrant infundibulogenesis integral to formulating a conceptual model for an infundibulocystic pathway to squamous cell carcinoma. Papillomatous and endophytic squamous proliferation is present with keratin plug, parakeratin, and mild pleomorphism of keratinocytes extending to the base of the biopsy (a). Clinically, these are solitary or coalescent skin-colored papules with a papillomatous surface. Given its papillomatous architecture and large size, many regarded giant condyloma acuminatum as a variant of verrucous carcinoma. Keratosis of unknown significance and leukoplakia: a preliminary study. 2015;19(2):198204. Multiple studies have shown that unlike giant condylomas, which consistently harbor HPV, anogenital verrucous carcinomas diagnosed on the basis of strict histopathologic criteria (see below) are predominantly negative for HPV.24,25,2830 These findings support distinct pathogenetic pathways in verrucous carcinoma and giant condyloma acuminatum, and that they should be considered separate entities. Hypertrophic Lichen Planus with Histological Features of Proliferative verrucous leukoplakia: a proposal for diagnostic criteria. Class 1 endophytes are mainly transmitted from host to host by vertical transmission, in which maternal plants pass fungi on to their offspring through seeds. Aside from koilocytosis, cytologic atypia is absent (Figure 2, B). Anyone you share the following link with will be able to read this content: Sorry, a shareable link is not currently available for this article. Mitotic activity is low and is confined to the lower third of the epithelium. Oral Surg Oral Med Oral Pathol Oral Radiol. As mentioned above, controversy exists regarding the relationship between verrucous carcinoma and giant condyloma acuminatum. PubMed Conspicuous fibrovascular cores and an infiltrative, rather than broad pushing, tumor base further distinguish papillary SCC from verrucous carcinoma. Biopsy of a lesion in the right proximal pretibial region demonstrated an atypical endophytic squamous proliferation, consistent with SCC. Among these three types of clavicipitaceous endophytes are different interactions with their plant hosts. These viral cytopathic changes serve as an important diagnostic hallmark (Figure 1, B).9 Binucleated or multinucleated forms may be present. This enhanced survivability is largely attributed to endophytic production of secondary metabolites which protect against herbivory as well as increased uptake of nutrients.
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