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By using our site, you agree to our collection of information through the use of cookies. To learn more, view our Privacy Policy. To browse Academia. Recent studies have suggested that the presence of endophytes in tall fescue can lead to decreased species richness in the associated plant community. To assess the generality of this hypothesis, a field study tested the effects of endophyte infection on a 3-yr-old successional field dominated by Festuca arundinacea. The potential importance of endophyte infection relative to other environmental factors was tested by including two additional treatments: the effects of soil fertility and mowing. Contrary to previous studies, a positive relationship was found between endophyte infection frequency and diversity , , ,. The relationship between endophytes and diversity varied throug The current COVID cruise tourism crisis has evolved to epic proportions and placed some of the cruise lines on the verge of bankruptcy. This research aimed to gain a deeper understanding of the crisis. Using an inductive qualitative approach, interviews were conducted with eight frequent cruisers who were at home and eight cruise ship employees who were employed by various cruise companies and who were working on cruise ships during the COVID cruise tourism crisis. The findings revealed a systematic failure within the cruise industry management to understand the COVID pandemic. Results of this study highlight the importance of health-related perceived risks on the nature and impact of the COVID cruise tourism crisis. This study supports the overall theory of cruise tourism and crisis management by extending the chaos theory and its principals on the COVID cruise tourism crisis. The managerial implications for cruise lines are outlined. Log in with Facebook Log in with Google. Remember me on this computer. Enter the email address you signed up with and we'll email you a reset link. Need an account? Click here to sign up. Variation in the abundance of fungal endophytes in fescue grasses along altitudinal and grazing gradients Dawn R Bazely. Related papers Broad-scale geographic patterns in the distribution of vertically-transmitted, asexual endophytes in four naturally-occurring grasses in Sweden Saewan Koh. A fungus among us: broad patterns of endophyte distribution in the grasses Jennifer Rudgers. A Sabedoria do Agora - Daniel J. Siegel Gelson Silva. Bazely John P. The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. Accepted 19 April Variation in the abundance of fungal endophytes in fescue grasses along altitudinal and grazing gradients Gustaf Granath, Mark Vicari, Dawn R. Bazely, John P. Granath Gustaf. Granath ebc. Present address of G. Vicari, D. Bazely, A. Puentes and T. Rakocevic, Biology Dept, York Univ. Present address of A. However, infection might also be associated with a cost to its host plant. Two differently-scaled approaches were used: 1 infection frequency was measured at a local scale along ten elevational transects within a ca km2 area and 2 infection frequency was measured on a regional scale along elevational transects on 17 mountains classified as having a history of high or low reindeer grazing pressure. Mean infection frequencies in F. Endophyte infection frequency in F. No elevational trends were observed in infection of F. These patterns in the distribution of endophyte-infected grasses in non-agricultural ecosystems may be explained by both biotic grazing and abiotic factors altitude. Differences in ecology and life history of the studied grass species may also be of importance for the different results observed among species. This is mainly due to the economic impact of endophyte-induced diseases on livestock productivity by means of toxic alkaloids Fribourg and Waller These alkaloids are also known to affect insect herbivores Clay and possibly wild vertebrate grazers Bazely et al. Fungal endophyte infection also has implications for plant community composition, because infected host grasses can grow more vigorously compared with uninfected plants Clay and Holah , Lewis For example, infection may increase drought resistance Elbersen and West , nutrient uptake Malinowski and Belesky and herbivore resistance Cheplick and Clay in host plants. If endophytes always benefit their host grass, then natural selection should always favour infected individuals, leading to high infection frequency in grass populations. This has been shown in several grazing studies Clay , Bazely et al. Knowledge about patterns of endophyte distribution in wild grasses is generally limited Brem and Leuchtmann , Faeth and Sullivan , Faeth et al. Thus, studies to investigate the variability of hostendophyte interactions in wild grass populations should be conducted in natural systems Saikkonen et al. In this study, we adopted the classical ecological approach of sampling along an ecotone to examine how endophyte distribution varies with abiotic and biotic factors in three widespread grass species: Festuca ovina , F. Endophyte infection in F. When E. The Scandinavian mountain range covers a wide range of altitudes and because there is variation in the movement of reindeer Rangifer tarandus herds, there are also herbivory gradients Moen and Danell Additionally, plant growth conditions generally become harsher with increasing altitude due to lower temperature, less favourable soil conditions, and shorter growing seasons Ericsson et al. A previous study of endophyte distribution in F. This observation is in line with previous findings that, under harsher conditions, endophytes may be less beneficial for their host grasses Ahlholm et al. An alternative hypothesis is that, in the Swedish mountains, herbivory may also be a key factor influencing the abundance and distribution of infected grasses, because endophytes could be functioning as an anti-herbivore defence Cheplick and Clay , Clay , Gwinn et al. If these general rules apply, we could expect that: A endophyte infection frequency in all three Festuca species decreases as altitude increases because the harsher conditions reduce the ability of the host grass to support its fungus. B endophyte infection frequency in all three Festuca species increases with increasing grazing pressure from reindeer since endophytes are providing an anti-herbivore defence by synthesizing a toxic alkaloid ergovaline. Methods Study areas Two scale-level approaches were employed to determine the abundance of endophyte infected grasses along altitudinal gradients. In the first, in August , we examined proportions of infected grass plants of the three fescue species in a mountain area near Abisko, Sweden Fig. Abisko is in the northern part of the Swedish mountains, and lies above the Arctic Circle. The tree line is between and m a. Locations of the study areas. A total of ten transects were located on a topographical map on nine mountains so as to cover a large altitudinal gradient within a ca km2 area. Festuca populations were patchily distributed along each transect, and tillers were sampled at specific sites wherever populations were encountered. Sites ranged in elevation from to m. Approximately 50 vegetative tillers were collected per site for each Festuca species present. If the population at the site covered a smaller area, the sampling interval between plants was reduced but was neverB2 m. Since mountains were sampled intensively at a local scale, it was not possible to categorize sites and transects with respect to the intensity of reindeer grazing pressure. The second approach was to determine gradients in levels of endophyte infection across a larger mountainous area, at a regional scale. In this part of the study, we sampled plants of the three grass species in August , from two regions further south of Abisko in the Swedish mountain range Fig. One elevational transect was sampled per mountain. In August , flowering culms or stems of Festuca plants were sampled instead of vegetative tillers. This is basically the same method performed by Saikkonen et al. However, research suggests that E. The benefits of sampling culms over vegetative tillers included the production of clearer, larger imprints on the immunoblot cards, and greater confidence in distinguishing F. Danell pers. Grazing pressure is estimated to have been stable over the last 30 yr, and likely for even longer. Reindeer grazing has had a large impact on plant community composition in the Swedish mountains: e. We considered reindeer to be the main vertebrate herbivore in our study, since summer grazing by microtine rodents has been found to have a low impact, even during peak population years Andersson and Jonasson In the B1 region we collected flowering culms from six mountains and in the B2 region we collected flowering culms from 11 mountains. The annual precipitation in the B1 region is mm and the tree line varies between and m a. Each station was sampled randomly within a circle of as large a diameter as possible up to a maximum of 30 m. The tillers and flowering stems were pulled up from the ground, wrapped in damp paper towels and placed in ziploc bags. Within 24 h, the stem was cut at the transition zone between the above-ground shoot and rhizome. Grass species Red fescue F. It occurs in diverse habitats: grasslands, meadows, moors, dunes, mountain slopes and wasteland. Above the tree line it is very often confined to moist lee-side habitats. It is slower-growing than F. It is generally tolerant to drought and low soil fertility, and can withstand close cutting and heavy grazing. Festuca vivipara is a common arctic species in northwestern Scandinavia, where it occurs in mountainous areas on dry, rocky, sandy ground or on peat soils. At higher altitudes it replaces, to some extent, the closely related F. Like F. Logistic regression Tabachnick and Fidell was used to confirm the linear regression analysis, but since the outcome of the two regression types gave identical results, only those from the linear regression are reported. These kits were developed for detection of Neotyphodium in perennial ryegrass Lolium perenne and tall fescue Festuca arundinacea in agricultural studies. However, Koh et al. Festuca rubra , Festuca ovina and Festuca pratensis. Approximately tillers were analyzed per immunoblot card, on which infected material leaves a red imprint. Ambiguous light pink imprints were confirmed by microscopic observation. Statistical analyses Statistical analyses were carried out with JMP v5. Indicator variable regression Kleinbaum and Kupper was used to test whether sample site altitude and grazing pressure predicted endophyte infection frequency in fescue populations sampled at sites along transects in area B regional scale. Since grazing pressure was not a variable in the more northerly Abisko study area local scale focus , simple linear regression was used to analyze these data. Residual analysis Tabachnick and Fidell was used to test whether the residuals were normally distributed. Since the number of tillers or flowering culms of each grass species varied at each site, the analyses were weighted by sample size Sokal and Rohlf Results Distribution of endophyte-infected grasses at the local scale In area A, around Abisko, the three Festuca species were patchily distributed along the ten elevational transects and did not always co-occur at each designated sample station. Consequently, the number of sample stations per transect per species varied from zero to six. Festuca ovina occurred along all ten transects, while Festuca rubra and Festuca vivipara were found along nine and five transects, respectively. At the two sites where a limited number of F. Endophyte infection was found in all three Festuca species, but infection frequency varied greatly among sample sites. The overall mean infection frequency for each species was similar Table 1. Since previous studies have often calculated the mean proportion of infection only for those sites where some infected grasses were present i. In the Abisko area, endophyte infection in F. This accounted for over half the variation Table 1. Overview of field sampling schemes and summary statistics for endophyte infection frequency in three Festuca grass species. Festuca vivipara is included for completeness in spite of the low sample size. The local scale study A was conducted north of the regional scale study B in the Swedish mountain range. Sampling scale area No. However, no altitudinal gradients in infection were observed in F. Festuca rubra and Festuca ovina were both sampled over a broad elevational range, while flowering plants of Festuca vivipara were only found in one area at higher altitudes B2. Flowering stems of F. These species co-occurred on 11 mountains. Festuca vivipara was sampled on 3 of the 11 mountains in area B2. Screening with immunoblot cards revealed a broad range of endophyte infection, but all five sample sites with F. Festuca rubra had significantly greater infection frequency than F. In order to facilitate comparison with previously published studies, infection frequencies in the subset of infected sites only was determined at: 1. For F. The proportion of infected plants increased with increasing altitude Table 3, Fig. The interaction i. The indicator variable regression for F. The number of sample sites where F. Discussion The levels of endophyte infection in the Swedish mountains were similar or lower to those observed in Finnish Festuca populations by Saikkonen et al. However, our results reveal, for the first time, that local and regional variation in endophyte infection in natural grasslands may be related to elevation and to herbivore grazing pressure. Mean infection in F. Infection frequencies in F. To our knowledge, this is the first report of endophyte infection in F. Table 2. Regression analysis of infection frequency versus altitude in the local scale study area A. Altitude was log10 transformed. Endophyte infection frequency proportions of infected plants at sample stations in Festuca ovina, F. Each point represents one sample station. The analysis was weighted by the number of vegetative tillers sampled at each site. The significant weighted linear regression analysis for F. The presence of significant elevational gradients, in which infection declined with increasing altitude in F. Consistent with this idea is the report by Ahlholm et al. However, when reindeer grazing levels were included as a variable at the regional level, the positive association between grazing pressure and infection frequency supported our second prediction: that the endophyte may provide an anti-herbivore defence mechanism, and thus it can be beneficial for its host grass Clay and Schardl Support for the endophyte-grass host interaction being mutualistic appears to hold in high nutrient agricultural systems Clay , Gwinn et al. Note however, that grazing by an unmanaged population of Soay sheep in natural grasslands on St. Kilda, Scotland was positively associated with endophyte infection in F. Our study highlights the challenges inherent in attempting to partition the effects of biotic herbivory and abiotic factors on endophyte distribution and abundance in unmanaged grasslands. This was especially so because the altitudinal trends in endophyte occurrence in F. Infection was not influenced by altitude at the local scale, but at the regional scale, and infection frequency in F. There are several factors that co-vary along altitudinal gradients that cannot be easily disentangled in this study. These factors can also vary depending on the ecology of a particular species, perhaps giving rise to the opposite trends that we observed. Festuca rubra is known to be less common at higher altitudes Danielsson , while F. Grazers Table 3. Results of indicator variable regression in Festuca ovina and F. Endophyte infection frequency proportions of infected plants at sample stations in Festuca ovina and F. The indicator regression analysis was weighted by the number of flowering culms sampled at each station. Grazing pressure was significant for F. Lines are associated with significant linear regressions for altitude p B0. Therefore, we could expect consumption of F. If this were the case, then the observed elevational trends in endophyte infection would be consistent with a positive response in infection frequency to herbivory pressure. Furthermore, the reindeer grazing pressure has been reported as very low in area A Olofsson et al. Festuca ovina would, therefore, be expected to support and retain its endophyte more effectively in harsh environments. It is interesting to ask whether the high infection frequency in F. Festuca vivipara plantlets, for example, have been shown to contain high nutrient content compared to seeds of F. Temperature is another important variable along an altitudinal gradient. The decline in infection frequency of F. This rhizomatous grass might outgrow its endophyte during prolonged cool periods when temperatures are high enough to support growth of the host but not the fungus. Festuca ovina and F. There are also genetic indications that E. Although our study cannot distinguish among all of those factors that vary with altitude, it highlights future research questions and hypotheses about the nature of endophyte-grass interactions in natural ecosystems. In particular, questions are raised about how the ecology and life-history of the host-grass, which is commonly overlooked, may be an important component in explaining variation in endophyte infection frequency. We thank them all. Thanks also to Joachim Strengbom for helpful comments on the manuscript. References Ahlholm, J. Vertically transmitted fungal endophytes: different responses of host-parasite systems to environmental conditions. Andersson, M. Rodent cycles in relation to food resources on an alpine heath. Bazely, D. Interactions between herbivores and endophyte-infected Festuca rubra from the Scottish islands of St. Kilda, Benbecula and Rum. Broad-scale geographic patterns in the distribution of vertically-transmitted, asexual endophytes in four naturally-occurring grasses in Sweden. Brem, D. Intraspecific competition of endophyte infected vs uninfected plants of two woodland grass species. Cheplick, G. Acquired chemical defences in grasses: the role of fungal endophytes. Effect of drought on the growth of Lolium perenne genotypes with and without fungal endophytes. Clay, K. Fungal endophytes of grasses: a defensive mutualism between plants and fungi. Interactions among fungal endophytes, grasses and herbivores. Fungal endophyte symbiosis and plant diversity in successional fields. Evolutionary origins and ecological consequences of endophyte symbiosis with grasses. Herbivore cause a rapid increase in hereditary symbiosis and alter plant community composition. Conover, M. Impact of the consumption of endophyte-infected perennial ryegrass by meadow voles. Danielsson, B. Ergovaline occurrence in grasses infected by fungal endophytes of semi-arid pastures in Spain. Food Agricult. Elbersen, H. Growth and water relations of field-grown tall fescue as influenced by drought and endophyte. Ericsson, G. Moose offspring body mass along an altitudinal gradient. Faeth, S. Are endophytic fungi defensive plant mutualists? Mutualistic asexual endophytes in a native grass are usually parasitic. Asexual Neotyphodium endophytes in a native grass reduce competitive abilities. Fribourg, H. Neotyphodium research and application in the USA. Blackwell, pp. Grime, J. Comparative plant ecology: a functional approach to common British species. Gwinn, K. Changes in Neotyphodium coenophialum infestation levels in tall fescue pastures due to different grazing pressures. Harmer, R. The growth and nutrient content of Festuca vivipara L. Hartley, S. Manipulation of nutrients and grazing levels on heather moorland: changes in Calluna dominance and consequences for community composition. Heggberget, T. Reindeer Rangifer tarandus and climate change: importance of winter forage. Jewell, P. Island survivors: the ecology of the Soay sheep of St Kilda. Ju, H. Temperature influences on endophyte growth in tall fescue. Kleinbaum, D. Applied regression analysis and other multivariable methods. Koh, S. Rapid detection of fungal endophytes in grasses for large-scale studies. Alpine plant life, 2nd ed. Leuchtmann, A. Lewis, G. Effects of biotic and abiotic stress on the growth of three genotypes of Lolium perenne with and without infection by the fungal endophyte Neotyphodium lolii. Malinowski, D. Adaptations of endophyte-infected cool-season grasses to environmental stresses: mechanisms of drought and mineral stress tolerance. Meijer, G. Reindeer in the Swedish mountains: an assessment of grazing impacts. Symbiosis between grasses and asexual fungal endophytes. Plant Biol. Olofsson, J. Importance of large and small mammalian herbivores for the plant community structure in the forest tundra ecotone. Saikkonen, K. Fungal endophytes: a continuum of interactions with host plants. Endophyte-grass-herbivore interactions: the case of Neotyphodium endophytes in Arizona fescue populations. Endophytic fungi in wild and cultivated grasses in Finland. Fungal endophytes: hitchhikers of the green world. Model systems in ecology: dissecting the endophyte-grass literature. Skogland, T. Wild reindeer foraging-niche organization. Skuncke, F. Renbeten och deras gradering. Sokal, R. Biometry: the principles and practice of statistics in biological research, 3rd ed. Tabachnick, B. Using multivariate statistics, 4th ed. Warenberg, K. Zabalgogeazcoa, I. Padre rico Padre pobre Daniel Viloria. Alexander, J. Suhu dan Kalornet Mansyur Nasution. De Nutellarum uaria natura Michel Aberson. Jurnal hukum Catcalling Reynaldi Rafi. Improbidade Administrativa Leonardo Primeiro. Mujer y empresa. Perspectivas y planteamientos Marcial Lopez. Effect of a 3-month L-carnitine supplementation and resistance training program on circulating markers and bone mineral density in postmenopausal women: a randomized controlled trial Emilia Samborowska. Thallophyta Herbivore.

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