(lolium perenne) and meadow fescue (festuca pratensis) - Poznań

University of Technology and Life Sciences in Bydgoszcz, Bydgoszcz, Poland 

OCCURRENCE AND ANTAGONISTIC PROPERTIES 

OF PERENNIAL RYEGRASS (LOLIUM PERENNE) 

AND MEADOW FESCUE (FESTUCA PRATENSIS) 

ENDOPHYTIC FUNGI 

D. Pańka and M. Jeske 

Abstract 

The objectives of the research were (i) to select perennial ryegrass and meadow 

fescue ecotypes colonized by Neotyphodium lolii and N. uncinatum respectively, (ii) to 

determine colonisation of selected ecotypes by other endophytic fungi, and (iii) to 

evaluate antagonistic properties of N. lolii and N. uncinatum towards chosen fungi in 

dual cultures assays. Neotyphodium spp. were endophytes most often isolated from 

perennial ryegrass and meadow fescue ecotypes. Acremonium spp. was the second 

most frequently isolated group of endophytes in meadow fescue. These fungi occurred 

in 52.3% of ecotypes. Acremonium spp. and Paecilomyces spp. were second 

most often detected fungi (21.4%) in perennial ryegrass ecotypes. The level of infection 

with other fungi was low and did not exceed 14.2% (Fusarium graminearum) 

in perennial ryegrass ecotypes and 23.8% (Phialophora-like sp.) in meadow fescue 

ecotypes. No antagonistic interaction between N. lolii, N. uncinatum isolates and 

endophytic fungi from the genus Acremonium and F. graminearum was detected. 

Growth inhibition zones were observed in combinations with Neotyphodium spp. 

and following test fungi: F. solani, F. culmorum and Phialophora-like sp. 

Key words: endophytes, Neotyphodium spp., perennial ryegrass, meadow fescue, 

ecotypes, dual culture 

Introduction 

Grasses have developed many symbiotic associations with fungi during their 

evolution. It includes both endo- and ectomycorrhizas and relationships in which 

fungi systemically infect grass shoots. Fungi that live their entire life inside the 

Phytopathologia 52: 29–40 

© The Polish Phytopathological Society, Poznań 2009 

ISSN 2081-1756

30 D. Pańka and M. Jeske 

over ground part of the host grass belong to the latter group. They are called 

endophytes. These fungi form nonpathogenic, systemic and usually intercellular 

associations (Bacon and DeBattista 1991). The most important and the best 

known grass endophytes belong to the Neotyphodium/Epichloë genera of the tribe 

Balansieae (Clavicipitaceae, Ascomycetes). They are often called e-endophytes. The 

most widely known fungi from this group are Neotyphodium lolii – the endophyte of 

perennial ryegrass, N. uncinatum – a symbiont of meadow fescue and N. coenophialum 

that colonizes tall fescue (Glenn et al. 1996). E-endophytes are not the only group 

of symptomless, seed-borne fungal symbionts of grasses. Two other discovered 

groups are called p-endophytes and a-endophytes. The former consists of closely 

related Gliocladium-like endophytes in perennial ryegrass and Phialophora-like 

endophytes in fescues (Latch et al. 1984, Philipson 1991, An et al. 1993). They are 

not related to e-endophytes and differ from them in many ways. The latter has been 

proposed to accommodate endophytes from the Acremonium genera (Naffaa et al. 

1998). Different endophytes may occur symbiotically in the same host plant (Latch 

et al. 1984, Philipson 1991). They interact with each other and can significantly influence 

the plant. Intensive research on Neotyphodium/Epichloë endophytes has allowed 

gathering ample information about associations they form. Effects of other 

endophytes on growth and development of the host plant are much less known. 

The aim of the study was to determine the occurrence of endophytic fungi in perennial 

ryegrass and meadow fescue ecotypes and their antagonistic properties. 

Materials and methods 

Perennial ryegrass and meadow fescue plants investigated in the study originated 

from various regions of Poland and were maintained in the ecotype collection 

at the Mochełek Research Station, University of Technology and Life Sciences, 

Bydgoszcz, Poland. Plants from the collection were screened for presence of 

Neotyphodium lolii and N. uncinatum in their tissues. The Phytoscreen Neotyphodium 

immunoblot kits (Agrinostics, Ltd. Co., Watkinsville, Georgia, USA) were used for 

this purpose. Cross sections of tiller bases were placed on a nitrocellulose membrane 

wetted with extraction buffer. After incubation overnight at 4°C, the membrane 

was blocked and monoclonal antibodies were added. After washing, second 

antibodies specific for the monoclonal antibodies were added. The membrane was 

washed again and protein-A with an alkaline phosphatase enzyme conjugate was 

added to complete the stacking effect. Then, the chromogen solution was added 

and a dark pink colour developed wherever the membrane bounded Neotyphodium 

spp. specific proteins (Phot. 1). 

Sixteen perennial ryegrass (LP) and 19 meadow fescue (FP) ecotypes infected 

with N. lolii and N. uncinatum, respectively, were chosen for further tests. Multiple 

isolations of endophytic fungi from plants were carried out. Three, 20 mm long, 

pieces of tiller base sampled from each ecotype were used for isolation of 

endophytic fungi in culture. The obtained pieces were divided into four cross sec-

Occurrence and antagonistic properties of perennial ryegrass... 31 

Phot. 1. Identification of Neotyphodium spp. in grass tillers using Agrinostics Phytoscreen 

immunoblot assay. Red marks on a nitrocellulose membrane indicate the endophyte presence 

(photo by D. Pańka) 

tions each. Surface sterilization was done by washing with 75% ethanol for 1 min 

and in 5% sodium hypochlorite for 1 min followed by three rinses in sterile distilled 

water for 2 min. Each part was then split further into a few small pieces 

which were placed on 9 cm Petri plates containing potato dextrose agar (PDA) medium 

(Difco) amended with 100 mg of penicillin G and 100 mg of dihydrostreptomycin 

sulphate to eliminate bacterial growth. Tissue segments were slightly 

pressed onto the surface of PDA medium. Petri plates were incubated at 22°C in 

darkness for 4–28 days. Fungi growing out from the inocula were transferred into 

fresh PDA medium, purified if needed and then transferred into PDA slants. Identification 

of endophytic fungal populations was performed on the basis of morphology 

of spores and fruiting bodies and cultural characteristics. Pathogenicity of 

some isolated fungi, potentially active as biological control agents (BCA), to perennial 

ryegrass and meadow fescue plants was checked. 

Antagonistic properties of N. lolii and N. uncinatum strains towards other 

endophytic or nonpathogenic fungi (test fungi): Acremonium sp., Fusarium solani, F. 

culmorum, F. graminearum and Phialophora-like sp. were tested in dual-culture assays 

in Petri plates with PDA medium. Three-week-old mycelial disks (5 mm of diameter) 

of N. lolii and N. uncinatum were placed on the surface of PDA in Petri plates 

about 3 cm from the edge of the plate and incubated for three–four weeks at 22°C 

in darkness (until colonies reached 10–15 mm of diameter). Then five 14-day-old 

mycelial discs of test fungi (5 mm of diameter) were placed in the Petri plates with 

N. lolii or N. uncinatum colony, keeping 2 cm distance between the fungal discs. 

There were four replicates for each combination. In control plates, only discs of the 

test fungi were placed centrally on PDA plate. Plates were incubated at 25°C in


darkness up to 28 days, depending on growth rate of the test fungi. Width of 

growth inhibition zone (Christensen 1996) between fungal cultures on a plate and 

individual biotic effect (Mańka 1974) for each assay were estimated. Measurements 

and observations were taken when colonies in control combinations 

reached the edge of the Petri plates. An average width of inhibition zone calculated 

from four replicates indicated the scale of inhibition in a combination. Two series 

of dual culture assays were performed. 

Results 

From each ecotype of perennial ryegrass and meadow fescue endophytic fungi 

from the genus Neotyphodium: N. lolii and N. uncinatum have been isolated respectively 

(Tables 1 and 2). Additionally, six species of other fungi colonizing perennial 

ryegrass have been obtained (Table 1). Second, most often isolated group of 

endophytes were Acremonium spp. and Paecilomyces spp. These fungi occurred in 

21.4% and 21.4% of the perennial ryegrass ecotypes, respectively. The perennial 

ryegrass infection with other fungi did not exceed 14.2%. Two Fusarium species: F. 

culmorum and F. graminearum were isolated from the perennial ryegrass ecotypes. 

Table 1 

Fungi isolated from perennial ryegrass (Lolium perenne) ecotypes 

Isolated fungi 

LP1 LP2 LP3 LP4 

Perennial ryegrass ecotypes 

LP5 LP6 LP7 LP8 LP9 LP10 LP11 LP12 LP13 LP14 

Neotyphodium lolii + + + + + + + + + + + + + + 

Acremonium spp. – – + + – – – – – – – – + – 

Fusarium culmorum – – – – – – – + – – – – – – 

Fusarium graminearum + – – – + – – – – – – – – – 

Paecilomyces spp. + – + + – – – – – – – – – – 

Colletotrichum 

graminicola 

– – + – – – – – – – – – – – 

Chaetomium spp. – – – – – – + – – – – – – – 

“+” – fungus presence in the ecotype, “ – ” – lack of the fungus in the ecotype. 

The level of meadow fescue infection with endophytic fungi was higher than that 

of perennial ryegrass (Table 2). Fungi from the genus Acremonium consisted the largest 

group (52.3%) of the isolated endophytes. The mycelium of Phialophora-like sp. 

was detected in five ecotypes of meadow fescue. Four Fusarium spp. were present: 

F. culmorum, F. graminearum, F. solani and F. poae but infection level of the meadow 

fescue ecotypes with these fungi did not exceed 9.5%. 

No antagonistic interactions between N. lolii, N. uncinatum isolates and the 

endophytic fungi from the genus Acremonium were detected (Table 3). Individual 

biotic effect in these combinations was neutral (value: 0). Therefore, there was no 

competition for space and nutrients between Neotyphodium spp. and Acremonium


Table 2 

Fungi isolated from meadow fescue (Festuca pratensis) ecotypes 

Meadow fescue ecotypes 

FP1 FP2 FP3 FP4 FP5 FP6 FP7 FP8 FP9 FP10 FP11 FP12 FP13 FP14 FP15 FP16 FP17 FP18 FP19 FP20 FP21 

Isolated fungi 

Neotyphodium uncinatum + + + + + + + + + + + + + + + + + + + + + 

Acremonium spp. + + + + – – – + – – + + – + – + – – + + – 

Phialophora-like sp. – – + – – – – – – + – – – + – – – – – + + 

Fusarium culmorum – – – + – – – – – – – – – – – – – – – – – 

Fusarium graminearum – – + – – – – – – – – – + – – – – – – – – 

Fusarium poae – – – – – – – – – – – – – – + – – – – – – 

Fusarium solani – – – + – – – – – – – – – – – – – – – – – 

Paecilomyces spp. + – – – – – – – – – – – – – – – – – – – – 

Gliomastix murorum – + – – – – – – – – – – – – – – – – – – – 

Chaetomium spp. – – – – – – – – – – + – – – – – – – – – – 

“+”–funguspresenceintheecotype,“–”–lackofthefungusintheecotype.


Table 3 

Antagonistic properties of Neotyphodium spp. isolates towards other fungi 

isolated from grasses – individual biotic effect of Neotyphodium spp. isolates 

Neotyphodium 

spp. isolate 

Acremonium spp. 

Fusarium 

solani 

Test fungi 

Fusarium 

culmorum 

Fusarium 

graminearum 

Phialophora-like 

sp. 

LP1 0 +1 

Perennial ryegrass 

+2 +4 +3 

LP2 0 +1 +2 +4 +3 

LP3 0 +1 +4 +4 +3 

LP4 0 +1 +4 +4 +3 

LP5 0 +1 +4 +3 +4 

LP6 0 +1 +4 +4 +3 

LP7 0 +2 +2 +4 +3 

LP8 0 +1 +4 +4 +4 

LP9 0 +2 +3 +4 +3 

LP10 0 +1 +3 +4 +3 

LP11 0 +2 +3 +3 +3 

LP12 0 +1 +2 +3 +3 

LP13 0 +1 +4 +4 +4 

LP14 0 +1 +3 

Meadow fescue 

+4 +3 

FP1 0 +2 +4 +4 +3 

FP2 0 +1 +4 +4 +3 

FP3 0 +1 +3 +4 +4 

FP4 0 +2 +4 +3 +3 

FP5 0 +1 +2 +4 +3 

FP6 0 +1 +3 +4 +3 

FP7 0 +1 +2 +4 +3 

FP8 0 +2 +4 +4 +4 

FP9 0 +1 +3 +3 +3 

FP10 0 +1 +3 +4 +3 

FP11 0 +1 +3 +4 +3 

FP12 0 +1 +4 +3 +2 

FP13 0 +1 +2 +3 +3 

FP14 0 +2 +4 +3 +4 

FP15 0 +1 +4 +4 +3 

FP16 0 +2 +3 +4 +3 

FP17 0 +1 +3 +4 +3 

FP18 0 +1 +3 +4 +3 

FP19 0 +1 +2 +4 +3 

FP20 0 +1 +2 +3 +3 

FP21 0 +1 +2 +4 +2


Table 4 

Effect of Neotyphodium spp. isolates on in vitro growth of other fungi isolated 

from grasses – average width of the inhibition zone between Neotyphodium spp. 

and test fungi (mm) 

Neotyphodium 

spp. isolate 

Acremonium spp. 

Fusarium 

solani 

Fusarium 

culmorum 

Test fungi 

Fusarium 

graminearum 

Phialophora-like 

sp. 

LP1 0 3 

Perennial ryegrass 

2 0 3 

LP2 0 4 2 0 3 

LP3 0 4 2 0 1 

LP4 0 2 2 0 3 

LP5 0 4 2 0 1 

LP6 0 4 1 0 3 

LP7 0 5 1 0 1 

LP8 0 4 2 0 2 

LP9 0 4 1 0 3 

LP10 0 3 3 0 2 

LP11 0 4 2 0 1 

LP12 0 2 2 0 2 

LP13 0 2 2 0 3 

LP14 0 4 2 

Meadow fescue 

0 3 

FP1 0 4 3 0 2 

FP2 0 4 2 0 3 

FP3 0 3 2 0 2 

FP4 0 4 2 0 3 

FP5 0 2 1 0 3 

FP6 0 4 1 0 3 

FP7 0 4 2 0 1 

FP8 0 5 2 0 2 

FP9 0 4 2 0 3 

FP10 0 4 3 0 3 

FP11 0 3 2 0 1 

FP12 0 4 2 0 3 

FP13 0 5 2 0 2 

FP14 0 4 1 0 2 

FP15 0 4 2 0 2 

FP16 0 4 2 0 3 

FP17 0 4 3 0 3 

FP18 0 3 3 0 3 

FP19 0 4 2 0 3 

FP20 0 3 2 0 3 

FP21 0 4 2 0 3


spp. The widest growth inhibition zones were measured in combinations with N. 

lolii, N. uncinatum and F. solani isolates (Table 4). LP7 and FP13 isolates were most 

effective in inhibiting the growth of F. solani. Development of Phialophora-like sp. 

was less inhibited. In most cases, the width of growth inhibition zone between colonies 

of this fungus and those of N. lolii and N. uncinatum reached 3 mm. In most 

cases, inhibition zones both for N. lolii and N. uncinatum combinations with F. 

culmorum did not exceed 2 mm. Growth of F. graminearum endophyte was not affected 

either by N. lolii or N. uncinatum isolates. Moreover, in combinations with 

this fungus the highest values of the individual biotic effect were noted. Lower values 

of this parameter were noted in combinations with F. culmorum and Phialophora-like sp. 

Discussion 

Endophytes – microorganisms living within plant tissues without causing visible 

symptoms have been found in almost all plants to date (Schulz et al. 1993). The 

role which some of them play inside a plant is still unknown but there is a huge 

group of endophytes affecting plants in different ways. They can change physiological 

activities of host plants influencing enhancements of biotic and abiotic stress 

(Carroll 1988, Hallmann and Sikora 1996, Sturz and Nowak 2000). Depending on 

plant species, different endophytes species are considered most important. The 

main symbionts of many grasses are fungi from Neotyphodium and Epichloë genera 

(Malinowski and Belesky 2000, Fribourg and Waller 2005, Zabalgogeazcoa and 

Bony 2005). In our study N. lolii and N. uncinatum were the most frequently isolated 

endophytic fungi from perennial ryegrass and meadow fescue ecotypes, respectively. 

High level of wild grasses infection with Neotyphodium spp. endophytes is 

well known and documented (Lewis et al. 1997, Faeth et al. 2001, Wäli et al. 2001, 

Pańka 2008). Low isolation level of these fungi occurs usually in commercial 

cultivars due to a long storage period under conditions unfavorable for endophytes 

(Rolston et al. 1986, Clay and Leuchtmann 1987, Cappelli and Buonaurio 2001, 

Pańka and Łukanowski 2001, Pańka and Sadowski 2002). 

Second most often isolated group of endophytic fungi in our experiments were 

Acremonium spp. in both grass species. This group of endophytes is not often isolated 

from grasses. They were found, for example, in Lolium multiflorum and Festuca 

paniculata, and are similar to Acremonium chilense – an endophyte of orchardgrass 

(Dactylis glomerata; Naffaa et al. 1996, 1998, Morgan-Jones et al. 1990). 

Gliocladium-like endophytes were not found in perennial ryegrass but Phialophora-like 

fungi were detected in some meadow fescue ecotypes tested in our study. An et al. 

(1993) detected p-endophytes in F. pratensis, F. gigantea and F. arizonica. In these 

grass species, these endophytes lived in cosymbiosis with Neotyphodium spp. 

endophytes. Moreover, serological analyses and PCR assay indicated that these 

p-endophytes and other two found in perennial ryegrass and tall fescue were 

closely related. In some reports p-endophytes are listed as relatively common 

cosymbionts of e-endophytes in many grass species (Latch et al. 1984, Gams et al.


1990, Philipson 1991, Siegel et al. 1995). On the contrary, Koga et al. (1994) did 

not detected Phialophora-like endophyte in meadow fescue ecotypes. Siegel and 

Latch (1991) reported that Phialophora-like endophyte from tall fescue showed an 

activity towards wide spectrum of pathogens in agar cultures. So, it may also have a 

positive effect on host fitness. Martyniuk (1986) observed higher resistance of cereals 

colonized with Phialophora spp. towards Gaeumannomyces graminis. This author 

isolated Phialophora spp. from grasses and cereals very often. 

Fungi other than Neotyphodium spp., Acremonium spp. and Phialophora-like were 

isolated only in a few cases in our experiment. Neotyphodium spp. as “strong” 

endophytes are usually present as the first ones inside a plant and could deter development 

of other fungi either by antagonistic activity or competitiveness for 

space and nutrients in a tiller base. This can be the reason for low detection level of 

these fungi. In our study, such antagonistic effect was observed in dual culture assays 

with test fungi: F. solani and F. culmorum. 

Neotyphodium lolii and N. uncinatum did not affect Acremonium spp. growth in dual 

culture assay. Moreover, the latter group of fungi was isolated relatively often. 

Therefore, the coexistence of different endophyte types can be probably found in 

many grass species in nature. The results obtained and literature data suggest that 

the positive effect observed in grasses infected with Neotyphodium spp. can be due 

not only to its contribution but to the presence in the same plant of other fungal 

endophytes and their synergistic activities in host protection (An et al. 1993). It is 

possible that, in many earlier tests of Neotyphodium benefits, it was unknown 

whether these endophytes were present alone in a plant or together with other 

endophytes. Therefore, it will be important to determine endophyte status of a 

plant prior an experiment in the future. 

Conclusions 

1. Perennial ryegrass and meadow fescue ecotypes tested in this study 

were commonly colonized with Neotyphodium lolii and N. uncinatum, respectively. 

2. Fungi from the genus Acremonium were the second most often isolated 

group of endophytes from meadow fescue ecotypes and Acremonium spp., 

Paecilomyces spp. – from perennial ryegrass ecotypes. 

3. Neotyphodium spp. did not show antagonistic activity towards Acremonium 

spp. or Fusarium graminearum in dual culture assay. 

4. Fungi from the genus Neotyphodium inhibited the growth of Fusarium 

solani, F. culmorum and Phialophora-like fungi in dual culture assay.


Streszczenie 

WYSTĘPOWANIE I WŁAŚCIWOŚCI ANTAGONISTYCZNE GRZYBÓW 

ENDOFITYCZNYCH ŻYCICY TRWAŁEJ (LOLIUM PERENNE) 

I KOSTRZEWY ŁĄKOWEJ (FESTUCA PRATENSIS) 

Celem badań było wyselekcjonowanie ekotypów życicy trwałej i kostrzewy 

łąkowej zasiedlonych przez, odpowiednio, Neotyphodium lolii i N. uncinatum oraz 

inne grzyby endofityczne oraz określenie właściwości antagonistycznych wyizolowanych 

endofitów rodzaju Neotyphodium w stosunku do wybranych grzybów. 

Najczęściej z badanych ekotypów życicy trwałej i kostrzewy łąkowej izolowano 

grzyby endofityczne rodzaju Neotyphodium (N. lolii i N. uncinatum). Drugą najczęściej 

izolowaną z kostrzewy łąkowej grupą endofitów były grzyby rodzaju Acremonium. 

Grzyby te występowały w 52,3% ekotypów. Acremonium spp. i Paecilomyces spp. były 

z kolei drugą co do wielkości grupą grzybów (21,4%) wykrywanych w ekotypach 

życicy trwałej. Poziom zasiedlenia przez inne gatunki grzybów był stosunkowo niski 

i nie przekraczał 14,2% (Fusarium graminearum) w ekotypach życicy trwałej i 

23,8% (gatunki podobne do Phialophora) w kostrzewie łąkowej. Nie zaobserwowano 

żadnych reakcji antagonistycznych między izolatami N. lolii i N. uncinatum aendofitami 

rodzaju Acremonium oraz F. graminearum. Strefy zahamowania wzrostu 

zanotowano na płytkach z Neotyphodium spp. i następującymi grzybami testowymi: 

F. solani, F. culmorum oraz gatunkami podobnymi do Phialophora. 

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Authors’ address: 

Dr. Dariusz Pańka, Dr. Małgorzata Jeske, Department of Phytopathology, 

University of Technology and Life Sciences in Bydgoszcz, ul. Kordeckiego 20, 

85-225 Bydgoszcz, Poland, e-mail: panka@utp.edu.pl 

Accepted for publication: 11.06.2009

(lolium perenne) and meadow fescue (festuca pratensis) - Poznań

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