Simulated folivory increases vertical transmission of fungal endophytes that deter herbivores and alter tolerance to herbivory in Poa autumnalis.

BACKGROUND AND AIMS: The processes that maintain variation in the prevalence of symbioses within host populations are not well understood. While the fitness benefits of symbiosis have clearly been shown to drive changes in symbiont prevalence, the rate of transmission has been less well studied. Many grasses host symbiotic fungi (Epichloë spp.), which can be transmitted vertically to seeds or horizontally via spores. These symbionts may protect plants against herbivores by producing alkaloids or by increasing tolerance to damage. Therefore, herbivory may be a key ecological factor that alters symbiont prevalence within host populations by affecting either symbiont benefits to host fitness or the symbiont transmission rate. Here, we addressed the following questions: Does symbiont presence modulate plant tolerance to herbivory? Does folivory increase symbiont vertical transmission to seeds or hyphal density in seedlings? Do plants with symbiont horizontal transmission have lower rates of vertical transmission than plants lacking horizontal transmission? METHODS: We studied the grass Poa autumnalis and its symbiotic fungi in the genus Epichloë. We measured plant fitness (survival, growth, reproduction) and symbiont transmission to seeds following simulated folivory in a 3-year common garden experiment and surveyed natural populations that varied in mode of symbiont transmission. KEY RESULTS: Poa autumnalis hosted two Epichloë taxa, an undescribed vertically transmitted Epichloë sp. PauTG-1 and E. typhina subsp. poae with both vertical and horizontal transmission. Simulated folivory reduced plant survival, but endophyte presence increased tolerance to damage and boosted fitness. Folivory increased vertical transmission and hyphal density within seedlings, suggesting induced protection for progeny of damaged plants. Across natural populations, the prevalence of vertical transmission did not correlate with symbiont prevalence or differ with mode of transmission. CONCLUSIONS: Herbivory not only mediated the reproductive fitness benefits of symbiosis, but also promoted symbiosis prevalence by increasing vertical transmission of the fungus to the next generation. Our results reveal a new mechanism by which herbivores could influence the prevalence of microbial symbionts in host populations.

Females engage in stronger relationships: positive and negative effects of shrubs are more intense for Poa ligularis females than for males.

Background and Aims: Dioecious plants are of particular concern in view of global environmental changes because reproductive females are more sensitive to abiotic stresses, thus compromising population viability. Positive interactions with other plants may counteract the direct effects of any abiotic environmental stress, allowing them to thrive and maintain a viable population in suboptimal habitats, although this process has not been tested for dioecious species. Furthermore, almost no data are available on the outcome of such species interactions and their link with local spatial patterns and sex ratios. Methods: We set up a field experiment with Poa ligularis, a dioecious native grass from the arid grasslands of South America. We studied the interaction of male and female plants with cushion shrubs of contrasting ecological strategies. We experimentally limited direct shrub-grass competition for soil moisture and transplanted plants to evaluate the amelioration of abiotic stress by shrub canopies (i.e. sun and wind) on grasses. We also studied the distribution of naturally established female and male plants to infer process-pattern relationships. Key Results: Positive canopy effects as well as negative below-ground effects were more intense for females than for males. Deep-rooted Mulinum spinosum shrubs strongly facilitated survival, growth and reproduction of P. ligularis females. Naturally established female plants tended to distribute more closely to Mulinum than co-occurring males. Female growth suffered intensive negative root competition from the shallow-rooted Senecio filaginoides shrub. Conclusions: Interactions with other plants may reduce or enhance the effect of abiotic stresses on the seemingly maladapted sex to arid environments. We found that these biased interactions are evident in the current organization of sexes in the field, confirming our experimental findings. Therefore, indirect effects of climate change on population sex ratios may be expected if benefactor species abundances are differentially affected.

Breeding system diversification and evolution in American Poa supersect. Homalopoa (Poaceae: Poeae: Poinae).

BACKGROUND AND AIMS: Poa subgenus Poa supersect. Homalopoa has diversified extensively in the Americas. Over half of the species in the supersection are diclinous; most of these are from the New World, while a few are from South-East Asia. Diclinism in Homalopoa can be divided into three main types: gynomonoecism, gynodioecism and dioecism. Here the sampling of species of New World Homalopoa is expanded to date its origin and diversification in North and South America and examine the evolution and origin of the breeding system diversity. METHODS: A total of 124 specimens were included in the matrix, of which 89 are species of Poa supersect. Homalopoa sections Acutifoliae, Anthochloa, Brizoides, Dasypoa, Dioicopoa, Dissanthelium, Homalopoa sensu lato (s.l.), Madropoa and Tovarochloa, and the informal Punapoa group. Bayesian and parsimony analyses were conducted on the data sets based on four markers: the nuclear ribosomal internal tanscribed spacer (ITS) and external transcribed spacer (ETS), and plastid trnT-L and trnL-F. Dating analyses were performed on a reduced Poa matrix and enlarged Poaceae outgroup to utilize fossils as calibration points. A relaxed Bayesian molecular clock method was used. KEY RESULTS: Hermaphroditism appears to be pleisiomorphic in the monophyletic Poa supersect. Homalopoa, which is suggested to have originated in Eurasia 8·4-4·2 million years ago (Mya). The ancestor of Poa supersect. Homalopoa radiated throughout the New World in the Late Miocene-Early Pliocene, with major lineages originating during the Pliocene to Pleistocene (5-2 Mya). Breeding systems are linked to geographic areas, showing an evolutionary pattern associated with different habitats. At least three major pathways from hermaphroditism to diclinism are inferred in New World Homalopoa: two leading to dioecism, one via gynodioecism in South America and another directly from hermaphroditism in North America, a result that needs to be checked with a broader sampling of diclinous species in North America. A third pathway leads from hermaphroditism to gynomonoecism in Andean species of South America, with strictly pistillate species evolving in the highest altitudes. CONCLUSIONS: Divergence dating provides a temporal context to the evolution of breeding systems in New World Poa supersect. Homalopoa The results are consistent with the infrageneric classification in part; monophyletic sections are confirmed, it is proposed to reclassify species of sect. Acutifoliae, Dasypoa and Homalopoa s.l. and it is acknowledged that revision of the infrageneric taxonomy of the gynomonoecious species is needed.

Abscisic acid metabolite profiling as indicators of plastic responses to drought in grasses from arid Patagonian Monte (Argentina).

The identification of hormonal and biochemical traits that play functional roles in the adaptation to drought is necessary for the conservation and planning of rangeland management. The aim of this study was to evaluate the effects of drought on i) the water content (WC) of different plant organs, ii) the endogenous level of abscisic acid (ABA) and metabolites (phaseic acid-PA, dihydrophaseic acid-DPA and abscisic acid conjugated with glucose ester-ABA-GE), iii) the total carotenoid concentration and iv) to compare the traits of two desert perennial grasses (Pappostipa speciosa and Poa ligularis) with contrasting morphological and functional drought resistance traits and life-history strategies. Both species were subjected to two levels of gravimetric soil moisture (the highest near field capacity during autumn-winter and the lowest corresponding to summer drought). Drought significantly increased the ABA and DPA levels in the green leaves of P. speciosa and P. ligularis. Drought decreased ABA in the roots of P. speciosa while it increased ABA in the roots of P. ligularis. P. ligularis had the highest ABA level and WC in green leaves. While P. speciosa had the highest DPA levels in leaves. In conclusion, we found the highest ABA level in the mesophytic species P. ligularis and the lowest ABA level in the xerophytic species P. speciosa, revealing that the ABA metabolite profile in each grass species is a plastic response to drought resistance.

Productividad en materia seca y captura de carbono en un sistema silvopastoril y un sistema tradicionalen cinco fincas ganaderas de piedemonte en el departamento de Casanare / Dry matter productivity and carbon sequestration in a silvopastoral System and a Traditional system in five foothill cattle farms in the department of Casanare

El objetivo fue comparar, en cuanto a productividad en materia seca y captura de carbono, entre un sistema silvopastoril y un sistema tradicional en cinco fincas ganaderas de piedemonte del departamento de Casanare. En el lote experimental de cada finca se sembró una mezcla de cuatro especies de pastos, de los cuales Brachiaria brizantha fue la especie que tuvo un desarrollo exitoso en las cinco fincas, y con la cual se analizó su productividad en materia seca y captura de carbono. A los cinco meses después de la renovación de pastos en los lotes experimentales se obtuvo una productividad en materia seca entre 6,3 y 14,9 tMS/ha, y encaptura de carbono entre 2,7 y 6,4 tCOT/ha; mientras que en los lotes tradicionales estuvo entre 4 y 5 tMS/ha, y en captura de carbono entre 1,7 y 2,9 tCOT/ha. A los 20 meses en los lotes experimentales se obtuvo una productividad en materia seca entre 12 y 13,7 tMS/ha, y en captura de carbono entre 4,1 y 4,5 tCOT/ha; mientras que en los lotes tradicionales estuvo entre 4,4 y 6,5 tMS/ha, y en captura de carbono entre 1,4 y 2,1 tCOT/ha. Esto indica que la renovación de praderas con pastos adecuados y el pastoreo de rotación favorecen la producción en materia seca y la captura de carbono...

The goal was to compare, in terms of dry matter productivity and carbon sequestration, asilvopastoral system with a traditional system in five foothill cattle farms in the Departmentof Casanare. A mixture of four species of grass was planted in the experimental plot of eachfarm, of which Brachiaria brizantha was the species that had a successful development in thefive farms, and whose dry matter and carbon sequestration productivity was analyzed. Fivemonths after pasture renovation in the experimental lots, a dry matter productivity between6.3 and 14.9 tMS/ha and carbon sequestration productivity between 2.7 and 6.4 tCOT/ha were obtained; in traditional lots, dry matter productivity was between 4 and 5 tMS/ha,while carbon sequestration productivity was between 1.7 and 2.9 tCOT/ha. At 20 months, dry matter productivity between 12 and 13.7 tMS/ha, and carbon sequestration productivitybetween 4.1 and 4.5 tCOT/ha were obtained in experimental lots, while in traditionallots, dry matter productivity was between 4.4 and 6.5 tMS/ha, while carbon sequestrationproductivity was between 1.4 and 2.1 tCOT/ha. This indicates that renewal of grasslandswith suitable pastures and rotational grazing favor dry matter and carbon sequestration production...

O objetivo foi comparar, com respeito à produtividade em matéria seca e captura de carbono,um sistema silvipastoril e um sistema tradicional em cinco fazendas de criação de gado nodepartamento de Casanare. No lote experimental de cada fazenda plantou-se uma misturade quatro espécies de pastos, dos quais Brachiaria brizantha foi a espécie que teve um desenvolvimentobem-sucedido nas cinco fazendas, e com a qual analisou-se sua produtividadeem matéria seca e captura de carbono. Cinco meses depois da renovação de pastos nos lotesexperimentais obteve-se uma produtividade em matéria seca entre 6,3 e 14,9 tMS/ha, e umacaptura de carbono entre 2,7 e 6,4 tCOT/ha; enquanto que nos lotes tradicionais esteveentre 4 e 5 tMS/ha, e uma captura de carbono entre 1,7 e 2,9 tCOT/ha. Aos 20 meses noslotes experimentais obteve-se uma produtividade em matéria seca entre 12 e 13,7 tMS/ha, eem captura de carbono entre 4,1 e 4,5 tCOT/ha; enquanto que nos lotes tradicionais esteveentre 4,4 e 6,5 tMS/ha, e uma captura de carbono entre 1,4 e 2,1 tCOT/ha. Isto indica quea renovação de pastos com variedades adequadas e o pastoreio de rotação favorecem a produçãode matéria seca e a captura de carbono...

Phenotypic plasticity as an index of drought tolerance in three Patagonian steppe grasses.

BACKGROUND AND AIMS: Despite general agreement regarding the adaptive importance of plasticity, evidence for the role of environmental resource availability in plants is scarce. In arid and semi-arid environments, the persistence and dominance of perennial species depends on their capacity to tolerate drought: tolerance could be given on one extreme by fixed traits and, on the other, by plastic traits. To understand drought tolerance of species it is necessary to know the plasticity of their water economy-related traits, i.e. the position in the fixed-plastic continuum. METHODS: Three conspicuous co-existing perennial grasses from a Patagonian steppe were grown under controlled conditions with four levels of steady-state water availability. Evaluated traits were divided into two groups. The first was associated with potential plant performance and correlated with fitness, and included above-ground biomass, total biomass, tillering and tiller density at harvest. The second group consisted of traits associated with mechanisms of plant adjustment to environmental changes and included root biomass, shoot/root ratio, tiller biomass, length of total elongated leaf, length of yellow tissue divided by time and final length divided by the time taken to reach final length. KEY RESULTS AND CONCLUSIONS: The most plastic species along this drought gradient was the most sensitive to drought, whereas the least plastic and slowest growing was the most tolerant. This negative relationship between tolerance and plasticity was true for fitness-related traits but was trait-dependent for underlying traits. Remarkably, the most tolerant species had the highest positive plasticity (i.e. opposite to the default response to stress) in an underlying trait, directly explaining its drought resistance: it increased absolute root biomass. The niche differentiation axis that allows the coexistence of species in this group of perennial dryland grasses, all limited by soil surface moisture, would be a functional one of fixed versus plastic responses.

Occurrence of the non-native annual bluegrass on the Antarctic mainland and its negative effects on native plants.

Few non-native species have colonized Antarctica, although increased human activity and accelerated climate change may increase their number, distributional range, and effects on native species on the continent. We searched 13 sites on the maritime Antarctic islands and 12 sites on the Antarctic Peninsula for annual bluegrass (Poa annua), a non-native flowering plant. We also evaluated the possible effects of competition between P. annua and 2 vascular plants native to Antarctica, Antarctic pearlwort (Colobanthus quitensis) and Antarctic hairgrass (Deschampsia antarctica). We grew the native species in experimental plots with and without annual bluegrass under conditions that mimicked the Antarctic environment. After 5 months, we measured photosynthetic performance on the basis of chlorophyll fluorescence and determined total biomass of both native species. We found individual specimens of annual bluegrass at 3 different sites on the Antarctic Peninsula during the 2007-2008 and 2009-2010 austral summers. The presence of bluegrass was associated with a statistically significant reduction in biomass of pearlwort and hairgrass, whereas the decrease in biomass of bluegrass was not statistically significant. Similarly, the presence of bluegrass significantly reduced the photosynthetic performance of the 2 native species. Sites where bluegrass occurred were close to major maritime routes of scientific expeditions and of tourist cruises to Antarctica. We believe that if current levels of human activity and regional warming persist, more non-native plant species are likely to colonize the Antarctic and may affect native species.

[The origin of polyploid genomes of bluegrasses Poa L. and gene flow between Northern Pacific and sub-Antarctic islands].

The involvement of present-day diploid bluegrass species in the formation of polyploid genomes was investigated using comparison of sequences of internal transcribed spacers ITS1 and ITS2, and the 5.8S rDNA sequence. It was demonstrated that highly polyploid New Zealand bluegrasses, P. cita (2n = 84; ca. 96 to 100), P. chathamica (2n = 112), and P. litorosa (2n = 263 to 266) formed separate highly supported clade together with tetraploids (2n = 28) P. intrusa, P. anceps, and P. trioides (Austrofestuca littoralis). Among the diploid species (2n = 14), the closest relatives of these species, as well as of the polyploid species of section Poa, are the genomes of Eurasian species P. remota, P. chaixcii (sect. Homalopoa), P densa (Bolbophorum), and P. sibirica (sect. Macropoa). Nuclear genomes of polyploid Stenopoa, Tichopoa, Oreinos, and Secundae are definitely related to the genome of Arctic species P. pseudabbreviata (sect. Abbreviatae). On the contrary, judging by the genes for nuclear 45S rRNA, genomes of diploid P. trivialis (sect. Pandemos), P. annua, and P. supina (sect. Ochlopoa both) are only remotely related to the genomes of highly polyploid species (distances p between them and other bluegrass species from different sections of subgenus Poa constitute 6-10% and 11-15%, respectively). The conclusion on the relationships between highly polyploid and diploid bluegrass species was tested using analysis of synapomorphic mutations in the 5.8S rRNA gene. It was demonstrated that genomes of Poa eminens (2n = 42) and P. schischkinii (2n = 70) (sect. Arctopoa both) were noticeably different in ITS regions from the genomes of the members of the type subgenus Poa. A comparison of the Arctopoa ITS regions showed that the differences between them constituted only 0.2%. At the same time, p distances between the Arctopoa ITS and those from the species belonging to other sections of the genus Poa varied from 5 to 14%. South American species P chonotica (sect. Andinae) (=Ncoraepoa chonotica) (2n = 42) was found to be related to Arctagrostis, Festucella, and Hookerochloa, being at the same time quite distant from the other species of the genus Poa. Polymorphic in chromosome number highly polyploid species of Northern Hemisphere, P. arctica (2n = 42 to 106), P. turneri (2n = 42, 63 to 64), and P. smirnovii (2n = 42, 70) (sect. Malacanthae) are relative to a large group of tetraploid (2n = 28) endemic bluegrass species from New Zealand and sub-Antarctic islands (P. novae-zelandiae and allied species).

Shifts in positive and negative plant interactions along a grazing intensity gradient.

Isolating the single effects and net balance of negative and positive species effects in complex interaction networks is a necessary step for understanding community dynamics. Facilitation and competition have both been found to operate in harsh environments, but their relative strength may be predicted to change along gradients of herbivory. Moreover, facilitation effects through habitat amelioration and protection from herbivory may act together determining the outcome of neighborhood plant-plant interactions. We tested the hypothesis that grazing pressure alters the balance of positive and negative interactions between palatable and unpalatable species by increasing the strength of positive indirect effects mediated by associational resistance to herbivory. We conducted a two-year factorial experiment in which distance (i.e., spatial association) from the nearest unpalatable neighbor (Stipa speciosa) and root competition were manipulated for two palatable grasses (Poa ligularis and Bromus pictus), at three levels of sheep grazing (none, moderate, and high) in a Patagonian steppe community. We found that grazing shifted the effect of Stipa on both palatable grasses, from negative (competition) in the absence of grazing to positive (facilitation) under increasing herbivore pressure. In ungrazed sites, belowground competition was the dominant interaction, as shown by a significant reduction in performance of palatable grasses transplanted near to Stipa tussocks. In grazed sites, biomass of palatable plants was greater near than far from Stipa regardless of competition treatment. Proximity to Stipa reduced the amount of herbivory suffered by palatable grasses, an indirect effect that was stronger under moderate than under intense grazing. Our results demonstrate that facilitation, resulting mainly from protection against herbivory, is the overriding effect produced by unpalatable neighbors on palatable grasses in this rangeland community. This finding challenges the common view that abiotic stress amelioration should be the predominant type of facilitation in arid environments and highlights the role of herbivory in modulating complex neighborhood plant interactions in grazing systems.

Nitrogen dynamics in the intact grasses Poa trivialis and Panicum maximum receiving contrasting supplies of nitrogen.

The C(3) grass Poa trivialis and the C(4) grass Panicum maximum were grown in sand culture and received a complete nutrient solution with nitrogen supplied as 1.5 mol m(-3) NH(4)NO(3). (15)N tracer techniques were used to quantify the relative use of root uptake and mobilization in supplying nitrogen to growing leaves in intact plants which either continued to receive nitrogen or which received the complete nutrient solution without nitrogen. The allocation of both (15)N-labelled nitrogen uptake and unlabelled mobilized nitrogen indicated that, under their conditions of growth, the sink strength of growing leaves was relatively greater in P. maximum than P. trivialis. The supply of nitrogen by mobilization to side tillers of P. trivialis was completely stopped as the external nitrogen supply was reduced, whilst in P. maximum some allocation of mobilized nitrogen to side tillers, roots and growing leaves was maintained. In both plant species receiving an uninterrupted supply of nitrogen the allocation pattern of mobilized nitrogen differed from that of nitrogen derived from root uptake. Differences exist in the degree to which P. trivialis and P. maximum utilized uptake and mobilization to supply nitrogen to the growing leaves. In P. trivialis roots were always a net sink of mobilized nitrogen, irrespective of the external nitrogen supply. In P. maximum, roots were a net sink of mobilized nitrogen when external nitrogen was withdrawn, but exhibited both source and sink behaviour when nitrogen supply was continued.