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On the origins of the tetraploid Bromus species (section Bromus, Poaceae): insights from internal transcribed spacer sequences of nuclear ribosomal DNA.

The internal transcribed spacer (ITS) region of nuclear ribosomal DNA from 22 diploid and tetraploid annual Bromus species of section Bromus (Poaceae) and three species belonging to other Bromus sections, Bromus catharticus (section Ceratochloa), Bromus anomalus (section Pnigma), and Bromus sterilis (section Genea), were investigated by PCR amplification and direct sequencing. The length of the ITS-1 region varied from 215 to 218 bp, and that of the ITS-2 region from 215 to 216 bp, in the species analyzed. ITS-1 was more variable and provided more informative sites (49) than ITS-2 (32). No variation was encountered within species. In pairwise comparison among species of section Bromus, sequence divergence ranged from 0.0 to 8.0% for the combined ITS-1 and ITS-2 regions. Parsimony analysis using Avena longiglumis and Hordeum vulgare as outgroups resulted in well-resolved phylogenetic trees and showed that section Bromus is monophyletic according to the species analyzed outside of the section. The analysis clarified the phylogenetic relationships among monogenomic (diploid) species. Introduction of the allotetraploid species did not change the general topology of the trees obtained using only the diploid species. Although some tetraploid-diploid species relationships will have to be clarified with faster evolving markers, the ITS sequences are shown to be useful for assessing evolutionary relationships among closely related Bromus species, as well as for clarifying taxonomic problems in previously controversial cases (e.g., Bromus alopecuros and Bromus caroli-henrici). New hypotheses are proposed concerning the origin of several allotetraploid species. For example, it is shown that the tetraploid Bromus hordeaceus diverged earlier than all other species of section Bromus, excluding the diploid B. caroli-henrici, which is found to be basal in this group. The tetraploid Bromus arenarius, which was considered a hybrid between sections Bromus and Genea, and the tetraploid Bromus adoensis are sister taxa within section Bromus; they belong in a weakly differentiated clade with the diploids Bromus brachystachys, Bromus japonicus, Bromus squarrosus, Bromus arvensis, and Bromus intermedius.

DNA, Plant↗

Bromus fasciculatus presl--a third diploid progenitor of Bromus section Genea allopolyploids (Poaceae).

Allozyme variation of ten heterozymes of seven enzymes among five accessions of a rare diploid Bromus fasciculatus was analysed with the use of PAGE and compared with that for six other species of section Genea of the genus Bromus. Allozymes charasteristic for diploids B. tectorum and B. fasciculatus are combined in fixed heterozygous phenotypes of tetraploid B. rubens. Fixed heterozygous phenotypes of tetraploid B. madritensis combine one allozyme of B. fasciculatus with another of diploid B. sterilis at each of the loci studied. Of the three diploids studied, only B. sterilis fits well for a role of a genome donor for the polyploid B. diandrus-rigidus complex. Bromus fasciculatus thus appears to be a diploid ancestor for the two tetraploids of section Genea, B. rubens and B. madritensis.

Bromus↗

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem.

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C3 and C4). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH4+ input in spring, whereas there were no differences for surface input of NO3-. Soil inorganic N was dominated by NH4+, but invaded sites had greater subsurface soil NO3-. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil delta15N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar delta15N values suggest that Bromus assimilated subsurface NO3-, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil delta15N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO3- concentrations were greater in the invaded communities during periods of ample precipitation, and NO3- leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO3- from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.

Biodiversity↗

Bromus species in winter wheat-population dynamics and competitiveness.

The infestation of Bromus species in small grains, especially in winter cereals has increased over recent years. In some areas winter wheat growers consider Bromus spp. as their worst grass weed. Besides yield reduction, Bromus spp. cause lodging and complication of harvest. In Germany the two most dominant species are Bromus sterilis (L.) and Bromus secalinus (L.). In order to develop control strategies the population dynamics of the weeds were investigated. Based on the results a deterministic mathematical model using differential and algebraic equations was used to estimate changes in the population of the two Bromus species.

Agriculture↗

[Cartography and geographical spread of the adventitious species of brome (Bromus spp.) among cereals in the Sais area of Morocco].

Bromus spp is causing serious problems in wheat in the Sais area. However, the damage of this weed varies from one region to another according to the agro-climatic conditions and crop systems. The characterization of the infestation level in each situation is a prerequisite to develop a control strategy adapted to each environment. This study was undertaken in order to determine the infestation level and geographical spread of the weedy brome (Bromus spp) on wheat in Sais following crop systems and pedo-climatic conditions. The results obtained during two consecutive years (1998-99 and 1999-2000) revealed that ripgut brome (Bromus rigidus Roth.) was the most dominant species in wheat fields in the surveyed regions, followed by B. rubens L., B. sterilis L., B. madritensis L. and B. mollus L. Among, 18 regions and 100 infested wheat fields, 16.67% of fields were slightly infested (Plant density of Bromus (Dbr < 90 plants/m2, 61.11% were moderately infested (90 < Dbr < 290 plants/m2) and 22.22% were highly infested (Dbr > 400 plants/m2). The maximum relative frequency was obtained with Bromus rigidus (47.15%) and the coverage was 40.43%. But, for B.rubens, B. madritensis and B. sterilis, the relative frequencies were respectively 31.42; 26 and 15% and their coverages were respectively 28.9, 20.4 and 12.5%.

Bromus↗

[Competitiveness of hard wheat (Triticum durum Desf.) varieties against ripgut brome (Bromus rigidus Roth)].

Varieties with an excellent competitiveness against ripgut brome (Bromus rigidus Roth.) would be very important to reinforce others methods to control ripgut brome weed. This study was carried out in 1999-2000 season in a greenhouse experiment to test the aggressiveness degree of six varieties of hard wheat (Oum Rabia, Isly, Marzak, Karim, Sebou, and Massa) combined with ripgut brome. Plant density was fixed at 16 plants of wheat or Bromus for pure crop and 8 plants for wheat and 8 for Bromus mixture. The results showed that the numbers of kernels/spikes were higher in the mixture for on pure composition. For the kernel weight, the result was opposite except for Isly and Marzak varieties. Karim and Isly varieties obtained the highest grain yield and were more competitive in mixture composition but Sebou and Massa varieties were less competitive against ripgut brome. Results of ripgut brome productivity and water use efficiency were similar and were used to determine the aggressiveness coefficient of hard wheat varieties against ripgut brome. The reduction of the shoot dry matter of brome was 22 to 56% at flowering. The grain yield of brome was reduced from 57 to 81%.

Bromus↗

[Evaluation of the seed stock and dynamics of ripgut brome (Bromus rigidus Roth) in the cultivation of wheat in the Sais area of Morocco].

The study was conducted on experimental plots differing in cultivation treatment (deep or shallow) at the National School of Agricultural station for three years. The objective was to study the influence of the tillage practices on vertical distribution of the seeds of ripgut brome (Bromus rigidus Roth) in the soil profile. Four tillage methods were tested: Zero tillage (SD), disc plough (PDL) and a disc harrow at two depths, 20 cm (CDM) and 30 cm (CDP). The results show that the type of cultivation affected the distribution of the seeds of ripgut brome. Shallow plots were highly infested by Bromus rigidus (876 plant/m2). The total number of seeds of ripgut brome was 96 and 45% in 0-4 cm of the soil, 3 and 29% in 4-15 cm and 1 and 26% in 15-30 cm, respectively of the deep (CDP) and shallow soil (SD). Deep tillage reduced the seed bank of ripgut brome by 96.6% and 66.7% for shallow. Seeds of ripgut brome survive more than three years at the deep soil. Processes of germination of this weed became functional after 98 days in the soil. The disc plough reduced significantly the brome population and seed bank compared to other tillage methods.

Agriculture↗

Capillary zone electrophoretic determination of phenolic compounds in chess (Bromus inermis L.) plant extracts.

A simple CZE method for quantification of phenolic compounds (vanillin, cinnamic, sinapic, chlorogenic, syringic, ferulic, benzoic, p-coumaric, vanillic, p-hydroxybenzoic, rosmarinic, caffeic, gallic and protocatechuic acids) in less than 10 min using 20 mM sodium tetraborate (pH 9.2) with 5% v/v methanol as a BGE and with UV detection at 254 nm is described. The LODs (3 S/N) ranged between 0.02 and 0.12 microg/ mL. Repeatabilities (RSDs) were 0.66-1.8 and 1.56-4.23% for migration times and peak areas (n = 5), respectively. The method was applied to the determination of phenolic compounds in chess (Bromus inermis L.) after Soxhlet extraction and purification of the crude extracts with SPE procedures. The results compared well with those obtained by liquid chromatographic method. B. inermis was found as a suitable model plant containing a broad spectrum of phenolic compounds in easily detectable concentrations and as a potential source of antioxidants.

Bromus↗

Do co-occurring plant species adapt to one another? The response of Bromus erectus to the presence of different Thymus vulgaris chemotypes.

Local modification of the soil environment by individual plants may affect the performance and composition of associated plant species. The aromatic plant Thymus vulgaris has the potential to modify the soil through leaching of water-soluble compounds from leaves and litter decomposition. In southern France, six different thyme chemotypes can be distinguished based on the dominant monoterpene in the essential oil, which is either phenolic or non-phenolic in structure. We examine how soils from within and away from thyme patches in sites dominated by either phenolic or non-phenolic chemotypes affect germination, growth and reproduction of the associated grass species Bromus erectus. To do so, we collected seeds of B. erectus from three phenolic and three non-phenolic sites. Seeds and seedlings were grown on soils from these sites in a reciprocal transplant type experiment in the glasshouse. Brome of non-phenolic origin performed significantly better on its home soil than on soil from a different non-phenolic or a phenolic site. This response to local chemotypes was only observed on soil collected directly underneath thyme plants and not on soil in the same site (<5 m away) but where no thyme plants were present. This is preliminary evidence that brome plants show an adaptive response to soil modifications mediated by the local thyme chemotypes. Reproductive effort was consistently higher in brome of phenolic origin than in brome of non-phenolic origin (on both thyme- and grass-soil), indicating that life-history variation may be related to environmental factors which also contribute to the spatial differentiation of thyme chemotypes. Moreover, we found that brome growing on thyme-soil in general was heavier than when growing on grass-soil, regardless of the origin of the brome plants. This is concordant with thyme-soil containing higher amounts of organic matter and nitrogen than grass-soil. Our results indicate that patterns of genetic differentiation and local adaptation may modify competitive interactions and possible facilitation effects in natural communities.

Adaptation, Physiological↗

Seasonal and spatial patterns of metals at a restored copper mine site II. Copper in riparian soils and Bromus carinatus shoots.

Soil and plants were sampled throughout winter and spring near a perennial stream traversing a restored mine site in a winter-rainy climate. Within 1m of an acidic reach of the stream, soil had pH 3-5 and 50-100 microg/g "bioavailable" copper (extractable with 0.01 M CaCl2). Soil 2-3 m from the stream had pH 5-8 and lower (less than 3 microg/g) bioavailable copper. "Oxide-bound" copper (extractable with 2N HCl) was 50-100 microg/g at most locations. Copper concentrations in the shoots of field-collected Bromus carinatus declined from 20 microg/g in winter to 2 microg/g in spring at all sampling sites. A similar temporal pattern was found in plants grown under controlled conditions. Thus B. carinatus has a developmental program for control of shoot copper concentration, causing a seasonally-varying pattern of copper phytoaccumulation over a large range of copper availability in the soil.

Biodegradation, Environmental↗

Identification and immunogold localization of a novel bromegrass (Bromus inermis Leyss) peroxisome channel protein induced by ABA, cold and drought stresses, and late embryogenesis.

A cDNA (BG-15) was isolated through differential screening of a cDNA library made from an ABA-treated bromegrass (Bromus inermis Leyss) suspension cell culture. The 819 bp pair cDNA encoded a 174 amino acid polypeptide with a calculated molecular mass of 18.08 kD and isolectric point of 7.50. The deduced amino acid sequences for the cDNA were 29.5% and 32.6% homologous to the known amino acid-selective channel proteins of the chloroplastic outer membrane in pea and barley, but were highly homologous (55.6% to 83.2%) to the putative membrane channel proteins from rice and Arabidopsis. Immunogold localization demonstrated that the channel protein encoded by this cDNA was present on the peroxisome membrane. High stringency southern analysis revealed that 1 to 2 copies of the peroxisomal channel protein (PCP) genes were present in the bromegrass genome. Northern and Western blots revealed that the PCP gene was responsive to both cold and drought stresses, and was rapidly induced by ABA (75 microM). The transcript of the PCP gene also accumulated during late embryogenesis, but declined rapidly during germination. Data taken together, responsiveness of the PCP to cold and drought stresses, and accumulation during late embryogenesis suggest this novel peroxisomal channel protein is associated with sugar and fatty acid metabolism through fatty acid import or succinate export from peroxisome during desiccation tolerance and energy metabolism.

Abscisic Acid↗

A lipid transfer protein gene BG-14 is differentially regulated by abiotic stress, ABA, anisomycin, and sphingosine in bromegrass (Bromus inermis).

The objective was to investigate the expression of a lipid transfer protein gene (LTP) both in bromegrass (Bromus inermis) cells and seedlings after exposure to abiotic stresses, abscisic acid (ABA), anisomycin, and sphingosine. A full-length cDNA clone BG-14 isolated from bromegrass suspension cell culture encodes a polypeptide of 124 amino acids with typical LTP characteristics, such as a conserved arrangement of cysteine residues. During active stages of cold acclimation LTP expression was up-regulated, whereas at the final stage of cold acclimation LTP transcript level declined to pre-acclimation level. A severe drought stress induced the LTP gene; yet, LTP expression doubled 3 d after re-hydration. Both temperature and heat shock duration influence LTP induction; however temperature is the primary factor. Treatment with NaCl stimulated accumulation of LTP mRNA within 15 min and the transcripts remained at elevated levels for the duration of the salinity stress. Most interestingly, Northern blots showed LTP was rapidly induced not only by ABA, but also by anisomycin and sphingosine in suspension cell cultures. Of the three chemicals, ABA induced the most rapid and highest response in LTP expression as well as highest freezing tolerance, whereas sphingosine was the least active for both LTP expression and freezing tolerance.

Abscisic Acid↗

Ecological genetics of vernalization response in Bromus tectorum L. (Poaceae).

BACKGROUND AND AIMS: Bromus tectorum (cheatgrass or downy brome) is an exotic annual grass that is dominant over large areas of former shrubland in western North America. To flower in time for seed production in early summer, B. tectorum plants generally require vernalization at winter temperatures, either as imbibed seeds or as established seedlings. METHODS: Variation in response to increasing periods of vernalization as seeds or seedlings for progeny of ten full-sib families from each of four B. tectorum populations from contrasting habitats was studied. KEY RESULTS: As vernalization was increased from 0 to 10 weeks, the proportion of plants flowering within 20 weeks increased, weeks to initiation of flowering decreased, and seed yield per plant increased, regardless of whether plants were vernalized as seeds or seedlings. Most of the variation was accounted for by differences among populations. Plants of the warm desert population flowered promptly even without vernalization, while those of the cold desert, foothill and montane populations showed incremental changes in response variables as a function of vernalization period. Populations differed in among-family variance, with the warm desert population generally showing the least variance and the cold desert population the most. Variation among populations and among families within populations decreased as vernalization period increased, whereas the non-genetic component of variance showed no such pattern. CONCLUSIONS: Variation in vernalization response was found to be adaptively significant and apparently represents the result of contrasting selection regimes on a range of founder genotypes.

Bromus↗

Interspecific control of non-symbiotic carbon partitioning in the rhizosphere of a grass-clover association: Bromus madritensis-Trifolium angustifolium.

Grass-legume interaction in the rhizosphere was investigated in a greenhouse experiment with two annual species, bromegrass Bromus madritensis (L.) and clover Trifolium angustifolium (L.) grown in mono and mixed cultures. Partitioning of below-ground carbon between roots, respiration, and soil was measured after separate 2 h-labelling of each species with 14CO2 followed by a 9 d chase period. At the time of labelling, clover nodules were not yet fixing N2. Bromegrass grew much faster than clover. Shoot biomass of bromegrass was greater in the presence of clover than in monoculture. By contrast, both shoot and root biomass of clover was less in the presence of bromegrass than in monoculture. Carbon assimilation during the period of labelling was proportional to shoot biomass and partitioning above and below-ground did not differ among treatments. Absolute amounts of labelled C allocated to rhizosphere respiration was more in bromegrass than in clover (respectively 1.38 mg C against 0.75 mg C in monoculture and 1.79 mg C and 0.63 mg C in mixed culture). However, when expressed as a percentage of below-ground C allocation, rhizosphere respiration was lower in bromegrass than in clover, respectively, 38% and 45% in monoculture. In mixed culture, this percentage increased by 7.3% for clover, and 3.5% for bromegrass, thus indicating that the interspecific effect of grass was higher than that of clover. The percentage of below-ground C in a soil solution of clover in mixed culture was more than 2-fold that measured in monoculture. It was also significantly correlated with the percentage of below-ground C in respiration. These results provided evidence that the grass-legume mixture has the potential to influence the rhizosphere processes of each species in more than an additive way and that the effect of the interaction was stronger on clover than on bromegrass. The possible implications of this in grass-legume competition are discussed.

Biomass↗

Widespread occurrence of Neotyphodium-like endophyte in populations of Bromus tomentellus Boiss. in Iran.

Neotyphodium species occur as endophytic fungi in cool-season grasses around the world. The beneficial aspects of grass-Neotyphodium associations have provoked researchers to look for a novel association in plant species where this symbiotum has not been reported. We surveyed Russian bromegrass (Bromus tomentellus Boiss.) accessions from a germplasm collection for the presence of Neotyphodium spp. fungi and determined levels of endophyte infection in B. tomentellus populations in native rangelands of Iran. Among 50 collected accessions, symbiotic fungi were detected in 45 accessions without any symptoms of choke disease on host plants. In culture medium, fast-growing endophytes appeared from seeds after 7-14 days. Plants grown from seed collections were 80-100% infected. Based on morphological characteristics and PCR analysis, we concluded that this fungus is a member of the Neotyphodium group of endophytic fungi. Lack of apparent toxicity to grazing animals suggests a place for endophyte-infected B. tomentellus in rangeland renovation, providing this infected grass exhibits increased tolerance to abiotic stresses.

Bromus↗

Interaction between the endophytic fungus Epichloe bromicola and the grass bromus erectus: effects of endophyte infection, fungal concentration and environment on grass growth and flowering

Epichloe bromicola is an endophytic fungal species that systemically and perennially colonizes intercellular spaces of leaf blades, leaf sheaths and culms of Bromus grass species. E. bromicola causes choke disease in B. erectus, suppressing maturation of most, if not all, host inflorescences. In an investigation of the interaction between fungus and host, we used a quantitative polymerase chain reaction technique to estimate the amount of fungal DNA, and thereby fungal concentration, in host plants. Fungal concentration was directly correlated with vegetative vigour of the plant, as measured by longest leaf length, number of tillers and vegetative above-ground biomass, suggesting that, during vegetative growth, the endophytic fungus is most beneficial for the plant when present in high concentrations. In contrast, the reproduction of the plant, as measured by the number of functional inflorescences, was inversely correlated with fungal concentration: the majority of infected plants, and all that were associated with high concentrations of fungi, were diseased. Thus, the benefit of endophyte infection for the plant is coupled with the disadvantages of infertility. Fungal concentration was shown to be at least in part genetically determined because fungal concentration differed significantly in different plant-endophyte genotype combinations (symbiotum). In a field experiment with normal and CO2-enriched environments, elevated CO2 levels favoured fungal reproductive vigour over host reproductive vigour, suggesting that these plant endophytes would be at a selective advantage in a corresponding environmental-change scenario. We conclude that a dynamic and complex relationship between fungal endophyte infection, fungal concentration, genotype and environment affects growth and fecundity of B. erectus and should contribute to the evolution of these plant-fungal interactions.

Journal Article↗

Effects of different supplements on splanchnic oxygen consumption and net fluxes of nutrients in sheep consuming bromegrass (Bromus inermis) hay ad libitum.

Fifteen sheep (53 kg), with catheters in a hepatic vein, the portal vein and a mesenteric vein and artery, were offered a bromegrass (Bromus inermis) hay (104 g crude protein (CP), 700 g neutral-detergent fibre and 65 g acid-detergent lignin/kg dry matter (DM) ad lib. with different supplements to determine the effects on net flux of oxygen and nutrients across the portal-drained viscera (PDV) and liver. The sheep were unsupplemented (Control) or received 5 g DM/kg body weight (BW) of ground maize (M), 7 g DM/kg BW of soya-bean hulls (H) or 0.73 g DM/kg BW of a mix of feedstuffs high in rumenundegradable protein (P). Apparent digestible energy (DE) intakes were 5.3, 10.4, 10.6 and 6.7 (SE 0.74) MJ/d and apparent digestible CP intakes were 37, 50, 79 and 68 (SE 4.3) g/d for Control, M, H and P treatments respectively. Splanchnic tissue oxygen consumption rates were 0.23, 0.32, 0.30 and 0.27 (SE 0.054) mol/h, and oxidative metabolism accounted for 0.46, 0.31, 0.33 and 0.47 (SE 0.051) of DE intakes for Control, M, H and P treatments respectively. Supplements increased (P < 0.05) release of alpha-amino nitrogen (AAN) by the PDV (4.2, 17.5, 19.6 and 18.1 mmol/h for Control, M, H and P treatments respectively). Splanchnic net flux of AAN was not affected by supplement treatments. Hepatic release of urea-N was increased (P < 0.05) by supplement treatments (27, 40, 46 and 44 mmol/h for Control, M, H and P respectively); the P treatment increased (P < 0.05) and the H treatment tended (P = 0.10) to increase splanchnic release of urea-N (7, 10, 20 and 27 mmol/h for Control, M, H and P treatments respectively). Net flux of glucose across the PDV was -4.6, 1.4, -5.6 and -7.2 (SE 1.65) mmol/h for Control, M, H and P treatments respectively. Hepatic glucose released averaged 23 (SE 2.0) mmol/h and was not affected by treatment. Treatments M and H increased (P < 0.05) PDV release of propionate compared with the Control treatment (4.5, 15.5, 16.8 and 7.7 mmol/h for Control, M, H and P treatments respectively). Release of acetate by the PDV was 43, 97, 118 and 67 (SE 23.9) mmol/h for Control, M, H and P treatments respectively. In summary, different supplements of low-quality grass did not increase the efficiency of N metabolism by splanchnic tissues. Treatment P had little effect on net flux across splanchnic tissues of glucose, L-lactate, beta-hydroxybutyrate and volatile fatty acids (VFA).(ABSTRACT TRUNCATED AT 400 WORDS)

Animal Feed↗

Mixed inoculation alters infection success of strains of the endophyte Epichloë bromicola on its grass host Bromus erectus.

Within-host competition in multiply infected hosts is considered an important component of host-parasite interactions, but experimental studies on the dynamics of multiple infections are still rare. We measured the infection frequencies of four strains of the fungal endophyte Epichloë bromicola on two genotypes of its host plant Bromus erectus after single- and double-strain inoculation. Double-strain inoculations resulted in fewer double, but more single, infections than expected on the basis of infection frequencies in single-strain inoculations. In most cases, only one of the two strains established an infection, and strains differed in their overall competitive ability. This pattern resembles the mutual exclusion scenarios in some theoretical models of parasite evolution. In addition, competitive ability varied with host genotype, which may represent a mechanism for the coexistence of strains in a population. Hence, considering the genetic variation in both host and parasite may be important for a better understanding of within-host dynamics and their role in epidemiology or (co)evolution.

Biological Evolution↗