Are tropical fungal endophytes hyperdiverse?

Fungal endophytes are ubiquitous fungi that inhabit healthy plant tissues without causing disease. Endophytes have been found in every plant species examined to date and may be important, but often overlooked, components of fungal biodiversity. In two sites in a lowland, moist tropical forest of central Panama, we quantified endophyte colonization patterns, richness, host preference, and spatial variation in healthy leaves of two co-occurring, understory tree species [ Heisteria concinna (Olacaceae) and Ouratea lucens (Ochnaceae)]. From 83 leaves, all of which were colonized by endophytes, we isolated 418 endophyte morphospecies (estimated 347 genetically distinct taxa), most of which were represented by only a single isolate (59%). Among morphospecies encountered in more than one leaf (nonsingletons), we found evidence of host preference and spatial heterogeneity using both morphospecies frequencies and presence/absence records. Based on these data, we postulate that tropical endophytes themselves may be hyperdiverse and suggest that extrapolative estimates that exclude them will markedly underestimate fungal species diversity.     

The mycorrhizal status of vascular epiphytes in Bale Mountains National Park,Ethiopia

Roots of 28 species of epiphytic vascular plants were collected on tree trunks and branches at six afromontane forest sites between 1700 and 3300 m above sea level in Bale Mountains National Park, Ethiopia. Seven of the 28 epiphyte species were colonized by vesicular-arbuscular mycorrhizal fungi (VAM). Mycorrhizal colonization only occurred at two of the six sites examined, at 2900 m and 3300 m, but more than one type of VAM endophyte was present in each case. Three facultative epiphytic species were all highly colonized by VAM on the forest floor, whereas roots from epiphytic habitats were weakly colonized. No correlations were found between VAM colonization, fine root diameter and root hair length, but VAM colonization and root hair abundance were negatively correlated. The lack of VAM colonization of potential, epiphytic host species at the majority of the sites examined points to the dispersal of VAM propagules as the factor limiting mycorrhizal colonization of epiphytic habitats. It is suggested that root systems of hemiepiphytic tree species serve as corridors between forest floor and tree trunks through which VAM may spread via hyphal growth.     

Fungal endophyte communities in two tropical forests of southern India: diversity and host affiliation

Fifteen tree species from a tropical dry thorn forest and fifteen tree species from a tropical dry deciduous forest in the Mudumalai Wildlife Sanctuary, Nilgiri Biosphere Reserve, southern India, were surveyed for their foliar endophyte communities during the dry and wet seasons. Surface sterilized leaf segments of uniform dimension were plated on nutrient agar and culturable endophytes growing from the segments were identified. Endophyte diversity was greater in the dry thorn forest than in the dry deciduous forest in the dry season. Although the isolation frequency of culturable endophytes increased for both forests during the wet season, the assemblages were represented not by any unique fungal species but by the commonly occurring ones. Furthermore, although individual leaves were densely colonized by endophytes, only a few species of endophytes colonized the whole leaves; and, only a few fungal species dominated the foliar endophytic communities and were common for both forests during both dry and wet seasons. Thus, even under wet conditions that favour dispersal and infection by fungi, the endophyte diversity increased only marginally, an indication that certain tropical forests are not hyperdiverse with reference to fungal endophytes. This should be considered when using culturable endophyte diversity as a surrogate for estimating global fungal diversity.     

Simulated acid rain affects birch leaf endophyte populations

Endophytes were frequently isolated from mountain birch (Betula pubescens var. tortuosa (Ledeb.) Nyman) leaves at a subarctic site where natural air pollution is low. We tested whether simulated acid rain had any influence on the occurrence of endophytes. Dry controls with only ambient rain and irrigated controls treated with spring water of pH 6 were compared with acid treatments at pH 3 and pH 4, prepared by adding both sulphuric and nitric acids. Treatments began in 1985 and leaf samples were taken twice during the summer of 1992. Leaves were surface sterilized, five leaf disks from each leaf placed on malt extract agar, and growing colonies were counted and identified. The most frequently isolated endophyte from birch leaves was a Fusicladium anamorph of Venturia sp. (88% of all the isolates in July and 75% of all the isolates in August), followed by a sterile mycelium and Melanconium sp. The number of endophytes isolated and the species number increased from July to August. Endophytes were most frequently isolated from the basal part of the midrib. The percentage of colonization by endophytes was similar in short and long shoots. More endophytes were isolated from leaves of branches taken at 1 m height than at 2 m height. The stronger acid rain treatment (pH 3) reduced by approximately 25% the number of isolated endophytes in August. Treatments did not have any effect on species composition of endophyte assemblages in birch leaves. Offprint requests to: Marjo L. Helander.     

Endophytic fungi reduce leaf-cutting ant damage to seedlings

Our study examines how the mutualism between Atta colombica leaf-cutting ants and their cultivated fungus is influenced by the presence of diverse foliar endophytic fungi (endophytes) at high densities in tropical leaf tissues. We conducted laboratory choice trials in which ant colonies chose between Cordia alliodora seedlings with high (E_high) or low (E_low) densities of endophytes. The E_high seedlings contained 5.5 times higher endophyte content and a greater diversity of fungal morphospecies than the E_low treatment, and endophyte content was not correlated with leaf toughness or thickness. Leaf-cutting ants cut over 2.5 times the leaf area from E_low relative to E_high seedlings and had a tendency to recruit more ants to E_low plants. Our findings suggest that leaf-cutting ants may incur costs from cutting and processing leaves with high endophyte loads, which could impact Neotropical forests by causing variable damage rates within plant communities.     

Fungal endophytes and phytochemistry of oak foliage: determinants of oviposition preference of leafminers?

Sedentary insect herbivores, such as gallformers and leafminers, are usually non-randomly distributed among and within host plants. Dispersion of these insects is largely a function of female oviposition choice. In field experiments and observations spanning two growing seasons, we tested the hypothesis that selective oviposition on individual leaves within trees by the dominant herbivore of Emory oak, the monophagous leaf-miner Cameraria sp. nov., is determined by the probability of colonization by endophytic fungi. These fungi are alleged to act as plant mutualists by deterring, killing, or inhibiting the growth of insect herbivores. We found that leaves selected by females for oviposition and paired, unmined leaves were equally likely to be colonized by fungal endophytes. Furthermore, condensed and hydrolyzable tannin levels, purported inhibitors of fungal infection, and protein content did not vary between leaves selected by females and unmined leaves, or between leaves with and without endophyte infections. We conclude that female Cameraria do not choose leaves within trees for oviposition on the basis of propensity for endophytic fungal infection or on phytochemical parameters that might indicate probability of future infections. At this spatial scale at least, fungal endophytes do not explain the highly aggregated distribution of Cameraria among leaves and associated costs in terms of increased larval mortality. Fungal endophytes may, nevertheless, affect leafminer dispersion and abundance at larger spatial scales, such as host plant populations or species. We did find, however, that the amount of mining activity on leaves is positively associated with increased colonization by fungal endophytes. We suggest that mining activity increases endophyte fungal infections by facilitating spore germination and hyphal penetration into the leaf or by altering leaf phytochemistry. The facilitation of endophyte colonization by leafmining activity coupled with the lack of predictability of endophyte infections based on leaf phytochemistry and almost 100% infectivity of all oak leaves during sporadic wet years may prevent female leafminers from discriminating leaves for oviposition on the basis of current or future levels of endophytes in leaves.     

Endophytic and epiphytic phyllosphere fungi of Camellia japonica: seasonal and leaf age-dependent variations

Seasonal and leaf age-dependent variations in the endophytic and epiphytic phyllosphere fungal assemblages of Camellia japonica were examined and compared. Live leaves of C. japonica were collected in four seasons (May, Aug, Nov, Feb), and fungi were isolated from healthy-looking leaves of 0, 1, 2 and 3 y old. The infection rate and total number of endophytic fungi increased May-Feb, and species richness of endophytes increased as leaves aged. In contrast the infection rate of epiphytic fungi was 100% for all leaf ages at every sampling date. The total number of epiphytic fungi isolated was greatest in May and lowest in Aug. The species richness of epiphytes did not differ significantly by season or leaf age. Eight fungal species were recorded as major phyllosphere fungi of C. japonica. Seasonal variations were detected for the frequencies of Colletotrichum gloeosporioides, C. acutatum, and epiphytes Pestalotiopsis sp.1, Aureobasidium pullulans, Phoma sp.1 and Ramichloridium sp., whereas the frequency of the endophyte Geniculosporium sp.1 varied with leaf age. The frequency of the epiphyte Cladosporium cladosporioides varied with both season and leaf age.     

Endophytic fungal communities in woody perennials of three tropical forest types of the Western Ghats,southern India

Fungal endophytes of tropical trees are expected to be exceptionally species rich as a consequence of high tree diversity in the tropics and the purported host restriction among the endophytes. Based on this premise, endophytes have been regarded as a focal group for estimating fungal numbers because their possible hyperdiverse nature would reflect significantly global fungal diversity. We present our consolidated ten-year work on 75 dicotyledonous tree hosts belonging to 33 families and growing in three different types of tropical forests of the NBR in the Western Ghats, southern India. We conclude that endophyte diversity in these forests is limited due to loose host affiliations among endophytes. Some endophytes have a wide host range and colonize taxonomically disparate hosts suggesting adaptations in them to counter a variety of defense chemicals in their hosts. Furthermore, such polyphagous endophytes dominate the endophyte assemblages of different tree hosts. Individual leaves may be densely colonized but only by a few endophyte species. It appears that the environment (the type of forest in this case) has a larger role in determining the endophyte assemblage of a plant host than the taxonomy of the host plant. Thus, different tropical plant communities have to be studied for their endophyte diversity to test the generalization that endophytes are hyperdiverse in the tropics, estimate their true species richness, and use them as a predictor group for more accurate assessment of global fungal diversity.     

Endophytic fungal diversity of 2 sand dune wild legumes from the southwest coast of India

Endophytic fungi of 3 age classes (seeds, seedlings, and mature plants) and 5 tissue classes (cotyledons, seed coats, roots, stems, and leaves) of coastal sand dune legumes Canavalia cathartica and Canavalia maritima were assessed by plating surface-sterilized segments on malt extract agar. Forty-six fungal taxa comprising 6 ascomycetes, 33 mitosporic fungi, 2 zygomycetes, and 5 sterile morphospecies were recovered. There was no significant difference in the colonization frequency of endophytes between plant species (p = 0.4098, Student's t test). Among the age classes, endophytic fungi colonized over 90% of seedlings and mature plants. Similarly, among tissue classes, endophytic fungi colonized over 90% of root, stem, and leaf segments. Diversity and richness of endophytic fungi were higher in C. cathartica than in C. maritima. Rarefaction curves revealed a "higher expected number of species" in mature plants of C. cathartica and seedlings of C. maritima, whereas it was highest in leaves of both plant species. The most dominant endophyte, Chaetomium globosum, colonized over 50% of the root, stem, and leaf segments of C. maritima and over 50% of the root segments of C. cathartica. The colonization frequency of C. globosum was found to be 5%-12.5% in seeds and increased up to 40%-64.4% in seedlings or mature plants. Halosarpheia sp. was the only marine fungus recovered among the endophytes.     

An endophyte‐rich diet increases ant predation on a specialist herbivorous insect

1. All plants form symbioses with microfungi, known as endophytes, which live within plant tissues. Numerous studies have documented endophyte–herbivore antagonism in grass systems, but plant–endophyte–insect interactions are highly variable for forbs and woody plants. 2. The net effect of endophytes on insect herbivory may be modified by their interactions with higher trophic levels, such as predators. Including these multitrophic dynamics may explain some of the variability among endophyte studies of non‐grass plants, which are currently based exclusively on bitrophic studies. 3. The abundance of natural foliar endophytes in a Neotropical vine was manipulated and beetles were fed high or low endophyte diets. Experimental assays assessed whether dietary endophyte load affected beetle growth, leaf consumption, and susceptibility to ant predation. 4. Beetles feeding on high‐ versus low‐endophyte plants had almost identical growth and leaf consumption rates. 5. In a field bioassay, however, it was discovered that feeding on an endophyte‐rich diet increased a beetle's odds of capture by predatory ants nine‐fold. 6. Endophytes could thus provide an indirect, enemy‐mediated form of plant defence that operates even against specialist herbivores. We argue that a multitrophic approach is necessary to untangle the potentially diverse types of endophyte defence among plants.     

植物内生菌定殖检测技术及其应用

自然环境中内生菌定殖于植物体内,对宿主植物产生多种有益效应,但是内生菌定殖情况难以检测,相关研究不够深入系统。目前在该领域使用较为广泛的检测技术包括：荧光标记、抗生素标记、荧光定量PCR和高通量测序等。内生菌通过孔隙伤口和降解细胞壁等方式侵染植物,通过种子垂直传递核心内生菌。对植物内生菌定殖的侵染方式、定殖方法和检测技术进行了归纳和整理,介绍了内生菌多种侵染和迁移途径,总结了目前内生菌定殖在生物防治、促进植物生长和污染修复等方面的功能,综述了多种检测方式在应用中的特点,以期为内生菌定殖植物的相关研究及其应用提供参考依据。     

Light converts endosymbiotic fungus to pathogen, influencing seedling survival and niche-space filling of a common tropical tree, Iriartea deltoidea

Pathogens are hypothesized to play an important role in the maintenance of tropical forest plant species richness. Notably, species richness may be promoted by incomplete filling of niche space due interactions of host populations with their pathogens. A potentially important group of pathogens are endophytic fungi, which asymptomatically colonize plants and are diverse and abundant in tropical ecosystems. Endophytes may alter competitive abilities of host individuals and improve host fitness under stress, but may also become pathogenic. Little is known of the impacts of endophytes on niche-space filling of their hosts.Here we evaluate how a widespread fungal endophyte infecting a common tropical palm influences its recruitment and survival in natural ecosystems, and whether this impact is modulated by the abiotic environment, potentially constraining host niche-space filling. Iriartea deltoidea dominates many wet lowland Neotropical forests. Diplodia mutila is a common asymptomatic endophyte in mature plants; however, it causes disease in some seedlings. We investigated the effects of light availability on D. mutila disease expression.We found I. deltoidea seedlings to preferentially occur under shady conditions. Correspondingly, we also found that high light triggers endophyte pathogenicity, while low light favors endosymbiotic development, constraining recruitment of endophyte-infested seedlings to shaded understory by reducing seedling survival in direct light. Pathogenicity of D. mutila under high light is proposed to result from light-induced production of H(2)O(2) by the fungus, triggering hypersensitivity, cell death, and tissue necrosis in the palm. This is the first study to demonstrate that endophytes respond to abiotic factors to influence plant distributions in natural ecosystems; and the first to identify light as a factor influencing where an endophyte is placed on the endosymbiont-pathogen continuum. Our findings show that pathogens can indeed constrain niche-space filling of otherwise successful tropical plant species, providing unoccupied niche space for other species.     

Endophytic fungi in forest trees: are they mutualists?

Forest trees form symbiotic associations with endophytic fungi which live inside healthy tissues as quiescent microthalli. All forest trees in temperate zones host endophytic fungi. The species diversity of endophyte communities can be high. Some tree species host more than 100 species in one tissue type, but communities are usually dominated by a few host-specific species. The endophyte communities in angiosperms are frequently dominated by species of Diaporthales and those in gymnosperms by species of Helotiales. Divergence of angiosperms and gymnosperms coincides with the divergence of the Diaporthales and the Helotiales in the late Carboniferous about 300 million years (Ma) ago, indicating that the Diaporthalean and Helotialean ancestors of tree endophytes had been associated, respectively, with angiosperms and gymnosperms since ≥300 Ma. Consequently, dominant tree endophytes have been evolving with their hosts for millions of years. High virulence of such endophytes can be excluded. Some are, however, opportunists and can cause disease after the host has been weakened by some other factor. Mutualism of tree endophytes is often assumed, but evidence is mostly circumstantial. The sheer impossibility of producing endophyte-free control trees impedes proof of mutualism. Some tree endophytes exhibit either a pathogenic or a putatively mutualistic behaviour depending on the situation. The lifestyle (mutualism, commensalism, parasitism) of most tree endophytes is, however, not known. They are just there in the tissue and resume growth at the onset of natural senescence of the host tissue on which they eventually sporulate. Density of colonization of conifer needles by endophytes increases with needle age. It is postulated that the needles die as soon as colonization density reaches a threshold value. Normally, the threshold is not reached before the onset of natural senescence. The threshold value may, however, be reached earlier under some adverse conditions, e.g. lack of light in dense stands. As a consequence, endophytes kill the needles prematurely. Needle endophytes could, thus, be useful to eliminate “parasitic” needle mass, i.e. needles which consume more than they produce.     

Endophytic fungi increase the processing rate of leaves by leaf‐cutting ants (Atta)

1. Fungal endophytes are microfungi that reside asymptomatically inside of leaf tissues, increasing in density and diversity through time after leaves flush. Previous studies have suggested that the presence of fungal endophytes in the harvest material of leaf‐cutting ants (Atta colombica, Guérin‐Méneville) may negatively affect the ants and their fungal cultivar. 2. In the present study, it was tested whether the presence and diversity of fungal endophytes affected the amount of time necessary for leaf‐cutter ants to cut, process, and plant leaf material in their fungal garden. It was found that ants took 30–43% longer to cut, carry, clean, and plant leaf tissue with high relative to low endophyte abundance, and that the ants responded similarly to leaf tissue with high or low endophyte diversity. 3. It was further investigated whether the fungal cultivars' colonisation rate was greater on leaf material without fungal endophytes. No difference in the ants' cultivar colonisation rate on leaf tissue with high or low endophyte abundance was observed.     

Isolation of endophytes from leaves of <Emphasis Type="Italic">Neolitsea sericea</Emphasis> in broadleaf and conifer stands

Endophytic fungi were isolated from leaves of Neolitsea sericea, a major lauraceous tree in the laurel forests of southern Kyushu, collected from the understory layer of broadleaf and conifer stands. Cytosphaera sp. and a species of Ascomycetes in leaf blade segments, plus a xylariaceous species and Phomopsis spp. in petiole segments, were isolated at relatively high frequency. In general, isolation frequencies of endophytes were higher in petiole than blade segments. In blade segments, patterns of endophyte isolation were quite different among stands, while relatively similar in petiole segments. Significant effects of sampling sites or canopy vegetation were rarely detected. the understory layer of laurel forests. Neolitsea sericea is an evergreen broadleaf lauraceous tree, widely distributed in areas of eastern Asia. In southern Kyushu, it is one of the most common trees, growing both as canopy and understory species. In this study, endophytes were isolated from the leaves of N. sericea growing in the understory layer of conifer and broadleaf forest stands to survey the endophytes of N. sericea leaves and to examine the effect of the canopy layer on endophytic mycobiota in understory plants.     

Diversity and dynamics of fungal endophytes in leaves, stems and roots of Stellera chamaejasme L. in northwestern China

This study was conducted to explore fungal endophyte communities inhabiting a toxic weed (Stellera chamaejasme L.) from meadows of northwestern China. The effects of plant tissue and growth stage on endophyte assemblages were characterized. Endophytes were recovered from 50 % of the samples, with a total of 714 isolates. 41 operational taxonomical units (OTUs) were identified, consisting of 40 OTUs belonging primarily to Ascomycota and 1 OTU belonging to Basidiomycota. Pleosporales and Hypocreales were the orders contributing the most species to the endophytic assemblages. The total colonization frequency and species richness of endophytic fungi were higher in roots than in leaves and stems. In addition, for the plant tissues, the structure of fungal communities differed significantly by growth stages of leaf emergence and dormancy; for the plant growth stages, the structure of fungal communities differed significantly by plant tissues. This study demonstrates that S. chamaejasme serves as a reservoir for a wide variety of fungal endophytes that can be isolated from various plant tissues.     

Shifts in Symbiotic Endophyte Communities of a Foundational Salt Marsh Grass following Oil Exposure from the Deepwater Horizon Oil Spill

Symbiotic associations can be disrupted by disturbance or by changing environmental conditions. Endophytes are fungal and bacterial symbionts of plants that can affect performance. As in more widely known symbioses, acute or chronic stressor exposure might trigger disassociation of endophytes from host plants. We tested this hypothesis by examining the effects of oil exposure following the Deepwater Horizon (DWH) oil spill on endophyte diversity and abundance in Spartina alterniflora – the foundational plant in northern Gulf coast salt marshes affected by the spill. We compared bacterial and fungal endophytes isolated from plants in reference areas to isolates from plants collected in areas with residual oil that has persisted for more than three years after the DWH spill. DNA sequence-based estimates showed that oil exposure shifted endophyte diversity and community structure. Plants from oiled areas exhibited near total loss of leaf fungal endophytes. Root fungal endophytes exhibited a more modest decline and little change was observed in endophytic bacterial diversity or abundance, though a shift towards hydrocarbon metabolizers was found in plants from oiled sites. These results show that plant-endophyte symbioses can be disrupted by stressor exposure, and indicate that symbiont community disassembly in marsh plants is an enduring outcome of the DWH spill.     

Diverse non-mycorrhizal fungal endophytes inhabiting an epiphytic,medicinal orchid (<Emphasis Type="Italic">Dendrobium nobile</Emphasis>): estimation and characterization

Although the terrestrial and temperate orchids–fungal biology have been largely explored, knowledge of tropical epiphytic orchids–fungus relationships, especially on the ecological roles imparted by non-mycorrhizal fungal endophytes, is less known. Exploitation of the endophytic fungal mycobiota residing in epiphytic orchid plants may be of great importance to further elucidate the fungal ecology in this special habitat as well as developing new approaches for orchid conversations. The composition of fungal endophytes associated with leaves, stems and roots of an epiphytic orchid (Dendrobium nobile), a famous Chinese traditional medicinal plant, was investigated. Microscopic imaging, culture-dependant method and molecular phylogeny were used to estimate their entity and diversity. Totally, there were 172 isolates, at least 14 fungal genera and 33 different morphospecies recovered from 288 samples. Ascomycetes, coelomycetes and hyphomycetes were three major fungal groups. There were higher overall colonization and isolation rates of endophytic fungi from leaves than from other tissues. Guignardia mangiferae was the dominant fungal species within leaves; while the endophytic Xylariaceae were frequently observed in all plant tissues; Colletotrichum, Phomopsis and Fusarium were also frequently observed. Phylogenetic analysis based on ITS gene revealed the high diversity of Xylariacea fungi and relatively diverse of non-Xylariacea fungi. Some potentially promising beneficial fungi such as Clonostachys rosea and Trichoderma chlorosporum were found in roots. This is the first report concerning above-ground and below-ground endophytic fungi community of an epiphytic medicinal orchid, suggesting the ubiquitous distribution of non-mycorrhizal fungal endophytes in orchid plants together with heterogeneity and tissue specificity of the endophyte assemblage. Possible physiological functions played by these fungal endophytes and their potential applications are also discussed briefly. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Enzymatic activity of endophytic fungi of six native seedling species from Doi Suthep-Pui National Park,Thailand

Endophytic fungi were isolated from the stems, petioles, midribs, and leaves of seedlings of six native tree species collected from Doi Suthep-Pui National Park, Thailand. Endophytes were isolated from all tissue samples investigated, and taxa included five ascomycetes, eight anamorphic taxa, and numerous sterile mycelia. Twenty-six strains were tested for their ability to produce cellulase, mannanase, proteinase, and xylanase. The ability to produce these enzymes was distributed amongst the strains tested. Rainforest seedlings supported a diverse array of endophytes that have a wide range of enzymatic activities. The implication of enzyme production in relation to lifestyle abilities of the endophytes is discussed.     

内生真菌感染对宿主羊草抗病性的影响

以感染内生真菌的天然禾草羊草为实验材料,通过体外纯培养条件下的内生真菌、感染内生真菌的离体叶片和在体叶片对3种病原菌的抑菌实验,以探讨内生真菌对宿主植物羊草在抗病性方面的贡献。结果表明:体外纯培养条件下,分离自羊草的内生真菌Epichlobromicola对新月弯孢(Curvularia lunata)、根腐离蠕孢(Bipolaris sorokiniana)和枝孢霉(Cladosporium sp.)这3种病原菌都具有抑制作用,抑菌率分别达56.22%,46.93%和45.15%,且内生真菌培养滤液可以有效抑制这3种病原菌的孢子萌发,平均萌发率分别为30.4%,15.7%和16.4%;宿主植物叶片在离体条件下,内生真菌感染可以有效降低羊草叶片受C.lunata和C.sp.侵染后的病斑数或病斑长度,但对B.sorokiniana不起作用,甚至提高了叶片的病斑数及病斑长度,而离体叶片提取液对不同病原菌均有不同程度的抑制作用;在体条件下,内生真菌均可以通过降低叶片病斑数来增强羊草植株对这3种病原菌的抗性。由此看来,内生真菌E.bromicola对宿主植物羊草在抗病原菌侵染方面有一定的增益作用。