Characterization and practical use of self-compatibility in outcrossing grass species

BackgroundSelf-incompatibility (SI) systems prevent self-fertilization in several species of Poaceae, many of which are economically important forage, bioenergy and turf grasses. Self-incompatibility ensures cross-pollination and genetic diversity but restricts the ability to fix useful genetic variation. In most inbred crops it is possible to develop high-performing homozygous parental lines by self-pollination, which then enables the creation of F₁ hybrid varieties with higher performance, a phenomenon known as heterosis. The inability to fully exploit heterosis in outcrossing grasses is partially responsible for lower levels of improvement in breeding programmes compared with inbred crops. However, SI can be overcome in forage grasses to create self-compatible populations. This is generating interest in understanding the genetical basis of self-compatibility (SC), its significance for reproductive strategies and its exploitation for crop improvement, especially in the context of F₁ hybrid breeding.ScopeWe review the literature on SI and SC in outcrossing grass species. We review the currently available genomic tools and approaches used to discover and characterize novel SC sources. We discuss opportunities barely explored for outcrossing grasses that SC facilitates. Specifically, we discuss strategies for wide SC introgression in the context of the Lolium–Festuca complex and the use of SC to develop immortalized mapping populations for the dissection of a wide range of agronomically important traits. The germplasm available is a valuable practical resource and will aid understanding the basis of inbreeding depression and hybrid vigour in key temperate forage grass species.ConclusionsA better understanding of the genetic control of additional SC loci offers new insight into SI systems, their evolutionary origins and their reproductive significance. Heterozygous outcrossing grass species that can be readily selfed facilitate studies of heterosis. Moreover, SC introduction into a range of grass species will enable heterosis to be exploited in innovative ways in genetic improvement programmes.     

Assessing changes to ecosystem structure and function following invasion by Spartina alterniflora and Phragmites australis: a meta-analysis

Biological invasions resulting from anthropogenic activities are one of the greatest threats to maintaining ecosystem functioning and native biodiversity. Invasions are especially problematic when the invading species behaves as an ecosystem engineer that is capable of transforming ecosystem structure, function, and community dynamics. Of particular concern is the spread of emergent wetland grasses whose root systems alter hydrology and structural stability of soils, modify ecosystem functions, and change community dynamics and species richness. To address the threats posed to ecosystems across the globe, management practices focus on the control and removal of invasive grasses. However, it remains unclear how severely invasive grasses alter ecosystem functions and whether alterations persist after invasive grass removal, limiting our ability to determine if management practices are truly sufficient to fully restore ecosystems. Here, we conducted a meta-analysis to quantify ecological alterations and the efficacy of management following the invasion of Spartina alterniflora and Phragmites australis, two common and pervasive invaders in coastal wetlands. Our results indicate that S. alterniflora and P. australis significantly alter measures of ecosystem functioning and organismal abundance. Invaded ecosystems had significant elevations in abiotic carbon and nitrogen fixation and uptake in areas with invasive grasses, with differential photosynthetic pathways of these two grass species further explaining carbon fluxes. Moreover, evidence from our analyses indicates that management practices may not adequately promote recovery from invasion, but more data are needed to fully assess management efficacy. We call for future studies to conduct pairwise comparisons between uninvaded, invaded, and managed systems and provide research priorities.

     

Within-generation and transgenerational plasticity in growth and regeneration of a subordinate annual grass in a rainfall experiment

Precipitation changes may induce shifts in plant species or life form dominance in ecosystems, making some previously subordinate species abundant. The plasticity of certain plant functional traits of these expanding subordinate species may be one possible mechanism behind their success. In this study, we tested if the subordinate winter annual grass Secale sylvestre shows plasticity in growth and reproduction in response to altered environment associated with field-scale rainfall manipulations (severe drought, moderate drought, and watering) in a semiarid grassland, and whether the maternal environment influences offspring germination or growth in a subsequent pot experiment. Compared to control plots, S. sylvestre plants grew 38% taller, and produced 32% more seeds in severe drought plots, while plants in watered plots were 17% shorter, and had 22% less seeds. Seed mass was greatest in severe drought plots. Plants growing in drought plots had offspring with enhanced juvenile shoot growth compared to the progeny whose mother plants grew in watered plots. These responses are most likely explained by the decreased cover of previously dominant perennial grasses in severe drought plots, which resulted in wetter soil compared to control and watered plots during the peak growth of S. sylvestre. We conclude that the plasticity of this subordinate annual species in response to changing environment may help to gain dominance with recurring droughts that suppress perennial grasses. Our results highlight that exploring both within-generation and transgenerational plasticity of subordinate species may lead to a better prediction of changes in plant species dominance under climate change.

     

Trait differences between grass species along a climatic gradient in South and North America

Question: Are trait differences between grasses along a gradient related to climatic variables and/or photosynthetic pathway? Location: Temperate grassland areas of South and North America. Methods: In a common garden experiment, we cultivated C₃ and C₄ grasses from grasslands under different climatic conditions, and we measured a set of 12 plant traits related to size and resource capture and utilization. We described (1) interspecific plant trait differences along a climatic gradient defined by the precipitation and temperature at the location where each species is dominant and (2) the association between those plant trait differences and the photosynthetic pathway of the species. Results: Trait differences between grasses were related to the precipitation at the area where each species is dominant, and to the photosynthetic pathway of the species. Leaf length, leaf width, plant height, leaf area per tiller, specific leaf area, leaf δ¹³C ratio, and nitrogen resorption efficiency increased while leaf dry matter content and nitrogen concentration in senesced leaves decreased as precipitation increased. A proportion of these changes along the gradient was related to the photosynthetic pathway because dominant grass species in cold areas with low precipitation are mainly C₃ and those from warm and wet areas are C₄. Conclusions: A previous worldwide analysis showed that traits of graminoid species measured in situ changed slightly along climatic gradients (< 10% variance explained). In contrast, under a common environment we observed that (1) grass traits changed strongly along a climatic gradient (30‐85% variance explained) and, (2) a proportion of those changes were related to the association between photosynthetic pathway of the species and precipitation.     

Feed intake and milk production in dairy cows fed different grass and legume species: a meta-analysis

The aim of this meta-analysis was to compare feed intake, milk production, milk composition and organic matter (OM) digestibility in dairy cows fed different grass and legume species. Data from the literature was collected and different data sets were made to compare families (grasses v. legumes, Data set 1), different legume species and grass family (Data set 2), and different grass and legume species (Data set 3+4). The first three data sets included diets where single species or family were fed as the sole forage, whereas the approach in the last data set differed by taking the proportion of single species in the forage part into account allowing diets consisting of both grasses and legumes to be included. The grass species included were perennial ryegrass, annual ryegrass, orchardgrass, timothy, meadow fescue, tall fescue and festulolium, and the legume species included were white clover, red clover, lucerne and birdsfoot trefoil. Overall, dry matter intake (DMI) and milk production were 1.3 and 1.6 kg/day higher, respectively, whereas milk protein and milk fat concentration were 0.5 and 1.4 g/kg lower, respectively, for legume-based diets compared with grass-based diets. When comparing individual legume species with grasses, only red clover resulted in a lower milk protein concentration than grasses. Cows fed white clover and birdsfoot trefoil yielded more milk than cows fed red clover and lucerne, probably caused by a higher OM digestibility of white clover and activity of condensed tannins in birdsfoot trefoil. None of the included grass species differed in DMI, milk production, milk composition or OM digestibility, indicating that different grass species have the same value for milk production, if OM digestibility is comparable. However, the comparison of different grass species relied on few observations, indicating that knowledge regarding feed intake and milk production potential of different grass species is scarce in the literature. In conclusion, different species within family similar in OM digestibility resulted in comparable DMI and milk production, but legumes increased both DMI and milk yield compared with grasses.     

Effects of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain

1. We examined the effect of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain in northern Australia. Macroinvertebrates were sampled from four grass communities: (1) para grass, (2) hymenachne (Hymenachne acutigluma), a native perennial; (3) rice (Oryza meridionalis), a native annual, and (4) areas where para grass had been sprayed with herbicide. 2. Macroinvertebrate richness, abundance and community similarity showed very few differences among the grass communities, particularly in the epiphytic habitat. Benthic invertebrates showed some differences among grasses, with lower richness and abundance and different community structure associated with hymenachne. Herbicide control of para grass had no apparent effect on benthic invertebrates but reduced the abundance of epiphytic invertebrates in the short term. 3. The results of this study indicate that para grass has very little impact on macroinvertebrate communities, despite the changes to macrophyte communities. This is probably because para grass has similar physical structure to the native grasses and because none of these grasses contribute directly to aquatic food webs. Control of para grass using herbicide has little impact on aquatic invertebrates. This suggests that predicting the impact of weed invasion in wetlands requires an understanding of both the functional properties of macrophytes and the habitat preferences of the macroinvertebrates.     

Tall fescue EST-SSR markers with transferability across several grass species

Tall fescue (Festuca arundinacea Schreb.) is a major cool season forage and turf grass in the temperate regions of the world. It is also a close relative of other important forage and turf grasses, including meadow fescue and the cultivated ryegrass species. Until now, no SSR markers have been developed from the tall fescue genome. We designed 157 EST-SSR primer pairs from tall fescue ESTs and tested them on 11 genotypes representing seven grass species. Nearly 92% of the primer pairs produced characteristic simple sequence repeat (SSR) bands in at least one species. A large proportion of the primer pairs produced clear reproducible bands in other grass species, with most success in the close taxonomic relatives of tall fescue. A high level of marker polymorphism was observed in the outcrossing species tall fescue and ryegrass (66%). The marker polymorphism in the self-pollinated species rice and wheat was low (43% and 38%, respectively). These SSR markers were useful in the evaluation of genetic relationships among the Festuca and Lolium species. Sequencing of selected PCR bands revealed that the nucleotide sequences of the forage grass genotypes were highly conserved. The two cereal species, particularly rice, had significantly different nucleotide sequences compared to the forage grasses. Our results indicate that the tall fescue EST-SSR markers are valuable genetic markers for the Festuca and Lolium genera. These are also potentially useful markers for comparative genomics among several grass species.Electronic Supplementary Material Supplementary material is available for this article at .     

Sonoran Desert Ecosystem transformation by a C4 grass without the grass/fire cycle

Aim Biological invasions facilitate ecosystem transformation by altering the structure and function, diversity, dominance and disturbance regimes. A classic case is the grass–fire cycle in which grass invasion increases the frequency, scale and/or intensity of wildfires and promotes the continued invasion of invasive grasses. Despite wide acceptance of the grass–fire cycle, questions linger about the relative roles that interspecific plant competition and fire play in ecosystem transformations. Location Sonoran Desert Arizona Upland of the Santa Catalina Mountains, Arizona, USA. Methods We measured species cover, density and saguaro (Carnegiea gigantea) size structure along gradients of Pennisetum ciliare invasion at 10 unburned/ungrazed P. ciliare patches. Regression models quantified differences in diversity, cover and density with respect to P. ciliare cover, and residence time and a Fisher’s exact test detected demographic changes in saguaro populations. Because P. ciliare may have initially invaded locations that were both more invasible and less diverse, we ran analyses with and without the plots in which initial infestations were located. Results Richness and diversity decreased with P. ciliare cover as did cover and density of most dominant species. Richness and diversity declined with increasing time since invasion, suggesting an ongoing transformation. The proportion of old‐to‐young Carnegiea gigantea was significantly lower in plots with dominant P. ciliare cover. Main conclusions Rich desert scrub (15–25 species per plot) was transformed into depauperate grassland (2–5 species per plot) within 20 years following P. ciliare invasion without changes to the fire regime. While the onset of a grass–fire cycle may drive ecosystem change in the later stages and larger scales of grass invasions of arid lands, competition by P. ciliare can drive small‐scale transformations earlier in the invasion. Linking competition‐induced transformation rates with spatially explicit models of spread may be necessary for predicting landscape‐level impacts on ecosystem processes in advance of a grass–fire cycle.     

Insect assemblages change along a gradient of invasion by a nonnative grass

Because invasions by nonnative plants alter the structure and composition of native plant communities, invasions can alter the function of ecosystems for animals that depend on plants for food and habitat. We quantified effects of an invasion by a nonnative grass on the insect community in grasslands of southeastern Arizona. We sampled insects on 54 1-ha plots established across a gradient of invasion by Lehmann lovegrass (Eragrostis lehmanniana Nees), a perennial species native to southern Africa. Between 2000 and 2004, we captured 94,209 insects representing 13 orders, 91 families, and 698 morphospecies during 2,997 trap nights. Richness of families, richness of morphospecies, and overall abundance of insects decreased as dominance of nonnative grass increased. With every 100 g/m² increase in biomass of nonnative grass, the average number of insect families decreased by 5%, morphospecies decreased by 6%, and overall abundance decreased by 14%. In areas dominated by nonnative grass, 2 of 8 orders and 6 of 27 families of insects were present less frequently and one family was present more frequently; 5 of 8 orders and 6 of 27 families of insects were less abundant and 3 families were more abundant than in areas dominated by native grasses. As a result, this plant invasion altered the structure of the insect community, which has consequences for animals at higher trophic levels and for ecosystem processes, including decomposition and pollination. Because complete eradication of nonnative plants might be possible only rarely, maintaining stands of native vegetation in invaded areas may be an important practical strategy to foster persistence of animals in grasslands invaded by nonnative plants.     

Can seed-borne endophytes promote grass invasion by reducing host dependence on mycorrhizas?

Symbiotic interactions between plants and microorganisms have recently become the focus of research on biological invasions. However, the interaction between different symbionts and their consequences in host-plant invasion have been seldom explored. Here, we propose that vertically transmitted fungal endophytes could reduce the dependency of invasive grasses on mycorrhizal fungi allowing host establishment in those environments where the specific mutualist may be not present. Through analyzing published studies on nine grass species, we evaluated the effect of seed-borne Epichloë endophytes on the relationship of invasive and non-invasive grasses with arbuscular mycorrhizal fungi (AMF), a symbiosis known to be fundamental for plant fitness and invasion success. The endophyte effect on AMF colonization differed between invasive and non-invasive grasses, reducing mycorrhization only on invasive species but with no impact on their biomass. These results allowed us to propose that Epichloë endophytes could reduce the dependency of host plants on the mutualism with AMF, promoting host grass establishment and subsequent invasion. Simultaneous interactions with different types of mutualists may have profound effects on the host-plant fitness facilitating its range expansion. Our findings suggest that some specific mutualistic fungi such as epichloid endophytes facilitate host invasion by reducing the requirements of the benefits derived from other mutualisms.     

Soil physical properties and carbon/nitrogen relationships in stable aggregates under legume and grass fallow

Short-season fallow with legumes and/or grasses can restore the soil organic C and nitrogen (N) and improve soil structure. In this study, we accessed the effects of 2-season legume and grass fallow on structural properties and C/N relationships in aggregates of a sandy loam soil. Two legumes (Calopogonium mucunoides and Centrosema pubescens), and two grasses (Guinea grass (Panicum maximum) and goose grass (Eleusine indica) were used. Results showed that Calopogonium and Centrosema increased soil total porosity and reduced soil bulk densities, while goose grass increased bulk density and reduced total porosity of the soils at 0–15 and 15–30?cm depths. Guinea grass significantly increased the saturated hydraulic conductivity (50.4?cm?h^?1) and water holding capacity of the soils. Aggregates, 4.75 to 0.5?mm were greater in Guinea grass and least in goose grass fallowed soils. Calopogonium increased macro-aggregates at 0–15?cm soils by 48%, and mean weight diameter (MWD) by 44%. Organic carbon in 0.5–0.25?mm and <0.25?mm aggregate sizes was higher in Guinea grass soils. Generally, grasses had 4-fold increases of C:N contents in dry aggregates. In conclusion, short-season fallow with Guinea grass, Calopogonium and Centrosema, increased soil C and N and protected them from losses in stable aggregates.     

呼伦贝尔草地野生禾本科牧草资源及资源评价

对呼伦贝尔草地野生禾本科牧草的属种组成、生长习性及饲用价值进行了研究。结果表明,呼伦贝尔草地现有禾本科牧草42属112种,这些禾本科牧草大多为多年生草本植物,适应典型草原和丘陵草甸草原生境,饲用价值高,适口性良好,是植被组成中的重要组成成分,生态利用前景广阔,是本地区重要的牧草资源。对其资源进行评价,为其合理开发利用提供理论依据。     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

Phenotypes of Two Generations of Sporobolus airoides Seedlings Derived from Acroptilon repens‐invaded and Non‐invaded Grass Populations

Although the ecological impacts of invasive species are well known, the evolutionary impacts on recipient native grass communities are not. We suggest that remnant native plants may provide desirable seed sources for restoration and native plant production. Native populations exposed to the selective pressures associated with exotic invasion may retain traits that increase their ability to coexist with invasive species. Two generations of Sporobolus airoides Torr. (Alkali sacaton) plants derived from lineages collected from within long‐term invaded areas of Acroptilon repens (L.) DC (Russian knapweed) and from adjacent non‐invaded areas were propagated in a greenhouse to evaluate generational changes in phenotypic traits from the production environment. Given the difference in invasion history of the two populations, we hypothesized that invaded and non‐invaded subpopulations would differ phenotypically. Phenotypic measurements revealed that invaded subpopulations had greater vegetative growth, whereas non‐invaded subpopulations had increased sexual reproduction. Phenotypic expression changed from the first to the second generation, predominantly in the invaded subpopulation. Generational phenotypic shifts are disadvantageous for native seed production which requires a standard product to sell commercially. However, phenotypic variation may improve field seed survival. This research demonstrates the potential value of targeting post‐invasion remnant grass populations for restoration.     

Assessment of interaction between sheep and poorly palatable grass: a key tool for grassland management and restoration

Invasion of poorly palatable grasses due to abandonment or improper grazing management decreases pastures feeding value and biodiversity. With the aim to control their spread, we assessed the relationship between sheep foraging behavior and changes in aboveground phytomass, leaf traits, and chemical features of the tall grass Brachypodium rupestre and evaluated the effects of a B. rupestre-based diet on epithelium keratinization of rumen. Our results demonstrated that sheep became less selective throughout the experimental trials and B. rupestre decreased its aboveground phytomass. Some leaf traits showed significant changes (LDMC, LA, and ADL were higher in ungrazed areas; leaf nitrogen content was higher in the grazed ones). In addition, we detected an increase of the degree of epithelium keratinization of sheep. Thus high grazing pressure can be used to control the spread of B. rupestre, but negative effects on animal welfare due to the increase of rumen keratinization might be expected.     

Impacts of silica-based defences in grasses on the feeding preferences of sheep

Grasses, which dominate many terrestrial ecosystems, sustain high densities of grazing mammals, so are of great economic and ecological importance. Traditionally, grasses are thought to be adapted to tolerate grazing rather than defend against it; however, silica deposited in the leaves of grasses has recently been shown to act as a feeding deterrent to invertebrate herbivores and small mammals. This study assesses whether silica is effective as a feeding deterrent to larger mammalian herbivores. We assess the impact of manipulated silica levels in five grass species on the feeding preferences of sheep both within and between grass species.Sheep feeding behaviour was driven by between-species differences in palatability. Hence, within a single species silica addition did not cause significant changes in feeding preference. However, there were significant differences in both the feeding preferences and bite rates between grass species, and these differences were much more marked when the grasses had been exposed to high levels of silica. The impacts that silica had on preference were least pronounced in palatable species (e.g. Poa annua) compared with less-palatable species (e.g. Brachypodium pinnatum and Festuca ovina). Sheep fed for longer, took more bites and had a higher bite rate on the grass species with the lowest leaf silica concentrations, namely P. annua.Sheep were less affected by silica defences than smaller, non-ruminant herbivores, but the changes in species preference rankings caused by silica suggest it may lead to changes in sward composition. Further, in species that are already relatively low in palatability, silica-induced reductions in bite rate could potentially reduce forage intake rates, with consequences for sheep performance.     

Exploiting the Brachypodium Tool Box in cereal and grass research

It is now a decade since Brachypodium distachyon (Brachypodium) was suggested as a model species for temperate grasses and cereals. Since then transformation protocols, large expressed sequence tag (EST) databases, tools for forward and reverse genetic screens, highly refined cytogenetic probes, germplasm collections and, recently, a complete genome sequence have been generated. In this review, we will describe the current status of the Brachypodium Tool Box and how it is beginning to be applied to study a range of biological traits. Further, as genomic analysis of larger cereals and forage grasses genomes are becoming easier, we will re-evaluate Brachypodium as a model species. We suggest that there remains an urgent need to employ reverse genetic and functional genomic approaches to identify the functionality of key genetic elements, which could be employed subsequently in plant breeding programmes; and a requirement for a Pooideae reference genome to aid assembling large pooid genomes. Brachypodium is an ideal system for functional genomic studies, because of its easy growth requirements, small physical stature, and rapid life cycle, coupled with the resources offered by the Brachypodium Tool Box.     

Global grass (Poaceae) success underpinned by traits facilitating colonization,persistence and habitat transformation

Poaceae (the grasses) is arguably the most successful plant family, in terms of its global occurrence in (almost) all ecosystems with angiosperms, its ecological dominance in many ecosystems, and high species richness. We suggest that the success of grasses is best understood in context of their capacity to colonize, persist, and transform environments (the “Viking syndrome”). This results from combining effective long‐distance dispersal, efficacious establishment biology, ecological flexibility, resilience to disturbance and the capacity to modify environments by changing the nature of fire and mammalian herbivory. We identify a diverse set of functional traits linked to dispersal, establishment and competitive abilities. Enhanced long‐distance dispersal is determined by anemochory, epizoochory and endozoochory and is facilitated via the spikelet (and especially the awned lemma) which functions as the dispersal unit. Establishment success could be a consequence of the precocious embryo and large starch reserves, which may underpin the extremely short generation times in grasses. Post‐establishment genetic bottlenecks may be mitigated by wind pollination and the widespread occurrence of polyploidy, in combination with gametic self‐incompatibility. The ecological competitiveness of grasses is corroborated by their dominance across the range of environmental extremes tolerated by angiosperms, facilitated by both C₃ and C₄ photosynthesis, well‐developed frost tolerance in several clades, and a sympodial growth form that enabled the evolution of both annual and long‐lived life forms. Finally, absence of investment in wood (except in bamboos), and the presence of persistent buds at or below ground level, provides tolerance of repeated defoliation (whether by fire, frost, drought or herbivores). Biotic modification of environments via feedbacks with herbivory or fire reinforce grass dominance leading to open ecosystems. Grasses can be both palatable and productive, fostering high biomass and diversity of mammalian herbivores. Many grasses have a suite of architectural and functional traits that facilitate frequent fire, including a tufted growth form, and tannin‐like substances in leaves which slow decomposition. We mapped these traits over the phylogeny of the Poales, spanning the grasses and their relatives, and demonstrated the accumulation of traits since monocots originated in the mid‐Cretaceous. Although the sympodial growth form is a monocot trait, tillering resulting in the tufted growth form most likely evolved within the grasses. Similarly, although an ovary apparently constructed of a single carpel evolved in the most recent grass ancestor, spikelets and the awned lemma dispersal units evolved within the grasses. Frost tolerance and C₄ photosynthesis evolved relatively late (late Palaeogene), and the last significant trait to evolve was probably the production of tannins, associated with pyrophytic savannas. This fits palaeobotanical data, suggesting several phases in the grass success story: from a late Cretaceous origin, to occasional tropical grassland patches in the later Palaeogene, to extensive C₃ grassy woodlands in the early–middle Miocene, to the dramatic expansion of the tropical C₄ grass savannas and grasslands in the Pliocene, and the C₃ steppe grasslands during the Pleistocene glacial periods. Modern grasslands depend heavily on strongly seasonal climates, making them sensitive to climate change.     

Refuge effect of an unpalatable forb on community structure and grass morphology in a temperate grassland

The role of unpalatable plant species as biological antiherbivore refuges for palatable species is well-documented at community level particularly in harsh environments. In productive sub-humid temperate grassland subjected to domestic grazing, we examined the protective effect of Eryngium horridum on plant community structure and floristic composition, and evaluated whether these changes impacted on a number of morphological traits of grasses, related to grazing resistance. We also investigated, for a palatable grass species (Stipa neesiana) the existence of morphological differences between protected and unprotected plants and if this eventual variation was either plastic or genetic. The study consisted of a field survey where we compared paired patches, with and without E. horridum, and a greenhouse experiment where we evaluated individuals of S. neesiana coming from both patch types over a 11 months period. Patches dominated by E. horridum had lower richness and cover of forbs than patches without the forb, and similar richness but greater cover of cool-season tussock palatable grasses, which suggests a protective role on the latter. Grasses in these patches also had longer blades and sheaths and lower specific leaf area. The morphological differences of S. neesiana individuals collected from both patch types disappeared after 11 months growth in a common environment which revealed significant phenotypic plasticity in this species. These results suggest the existence of plant-to-plant facilitation in a productive ecosystem not only at community level, through changes in species richness and the promotion of palatable grasses, but also at population level, through plastic changes in aboveground morphological traits. Both facilitation and plasticity, would contribute to the persistence of threatened palatable grasses in the heavy grazed productive ecosystems.     

赖草属植物的分类现状及主要存在的问题

赖草属(Leymus Hochst.)为禾本科(Poaceae)小麦族(Triticeae)中的一个有重要经济价值的属,属内多数种类是优良牧草,有些种类具有耐寒、耐旱、耐碱等特性,是农业良种繁育、畜牧业牧草改良利用的重要基因资源。该属在分类学上是一个疑难属,在属的界限、属下组系的划分,以及类群间演化关系上问题较多。对赖草属分类学问题进行了综述,为赖草属植物资源的开发利用和保护提供资料。