OIKOS 98: 37–46, 2002
Infection by the systemic fungus Epichloe ¨ glyceriae and clonal
growth of its host grass Glyceria striata
Jean J. Pan and Keith Clay
Pan, J. J. and Clay, K. 2002. Infection by the systemic fungus Epichloe ¨ glyceriae and
clonal growth of its host grass Glyceria striata. – Oikos 98: 37–46.
Many plant species have the ability to expand laterally through space by clonal
growth. Plant pathogens can affect clonal growth characteristics thereby altering the
success of host plants within populations and of clonal species within communities.
We conducted a greenhouse experiment to determine the effects of the systemic
fungal pathogen, Epichloe ¨ glyceriae, on clonal growth patterns of its host grass,
Glyceria striata. We found that infected and uninfected plants produced similar total
biomass and numbers of tillers plus primary stolons per mother ramet. However,
biomass allocation to tillering (vegetative growth) vs stolon production (clonal
growth) was significantly affected by pathogen infection. Infected plants produced
more stolons and clonal growth biomass than uninfected plants while mother ramets
of uninfected plants produced more tillers and biomass than infected plants. Stolon
architecture of infected and uninfected plants also differed. In two of three popula-
tions, infected plants produced stolons with greater biomass and shorter spacer
lengths, even though mean stolon lengths were similar for infected and uninfected
plants. These results contrast strongly with most other clonal plant-pathogen systems
where infected plants are less vigorous and have reduced clonal growth compared to
uninfected plants. Greater clonal growth may be an effective mechanism for host
genotypes to persist and spread when seed production is prevented, as is the case with
castrating pathogens like Epichloe ¨ glyceriae.
J. J. Pan and K. Clay, Dept of Biology, Jordan Hall 142, 1001 E. Third St., Indiana
Uni., Bloomington, IN 47405, USA ( jepan@indiana.edu).
Clonal plant species are ubiquitous and important com-
ponents of many plant communities (Oborny and
Bartha 1995, Klimes et al. 1997), especially those dom-
inated by monocots, as approximately 70% of extant
monocot families are rhizomatous (Tiffney and Niklas
1985). Clonal plants commonly spread through both
seed production and clonal growth via rhizomes,
stolons, etc. (see Klimes et al. 1997 for a detailed
categorization of clonal growth). The ability to grow
clonally plays an important role in the persistence and
spread of clonal plant species (Janzen 1977, Cook 1979,
Eriksson and Jerling 1990, Oborny and Cain 1997), e.g.
the success of many weeds has been attributed to clonal
growth (Thompson 1991, Pys ˇek 1997). Clonal growth
also plays an important role at the population level.
Differences in clonal growth can affect competitive
interactions and lead to dominance by a few genotypes
(McLellan et al. 1997). Harberd (1961) found that one
genotype of the rhizomatous Festuca rubra had spread
about 220 meters laterally and estimated its age to be
about 400 years. Thus, changes to clonal growth char-
acteristics may alter the success of different plant geno-
types within populations.
Pathogens often modify host plant growth and repro-
duction (Burdon 1987, Isaac 1992). Infected host plants
generally have reduced fitness as a result of changes in
physiology, destruction of host tissue, infection of re-
productive structures, and are typically stunted and less
vigorous than their uninfected counterparts (Burdon
1987, Jarosz and Davelos 1995). Grasses infected by
systemic fungal endophytes commonly experience two
types of modifications to their growth: reproductive
Accepted 21 December 2001
Copyright © OIKOS 2002
ISSN 0030-1299
OIKOS 98:1 (2002) 37