Delayed induced silica defences in grasses and their potential for destabilising herbivore population dynamics

Some grass species mount a defensive response to grazing by increasing their rate of uptake of silica from the soil and depositing it as abrasive granules in their leaves. Increased plant silica levels reduce food quality for herbivores that feed on these grasses. Here we provide empirical evidence that a principal food species of an herbivorous rodent exhibits a delayed defensive response to grazing by increasing silica concentrations, and present theoretical modelling that predicts that such a response alone could lead to the population cycles observed in some herbivore populations. Experiments performed under greenhouse conditions revealed that the rate of deposition of silica defences in the grass Deschampsia caespitosa is a time-lagged, nonlinear function of grazing intensity and that, upon cessation of grazing, these defences take around one?year to decay to within 5?% of control levels. Simple coupled grass-herbivore population models incorporating this functional response, and parameterised with empirical data, consistently predict population cycles for a wide range of realistic parameter values for a (Microtus) vole-grass system. Our results support the hypothesis that induced silica defences have the potential to strongly affect the population dynamics of their herbivores. Specifically, the feedback response we observed could be a driving mechanism behind the observed population cycles in graminivorous herbivores in cases where grazing levels in the field become sufficiently large and sustained to trigger an induced silica defence response.     

Herbivore specific induction of silica-based plant defences

Induced plant responses to herbivory have major impacts on herbivore feeding behaviour, performance and population dynamics. These effects are well established for chemical defences, but induction of physical defences remains far less studied. However, for many plants, it is physical defences that play the major role in regulating the levels of herbivore damage sustained. We provide evidence that, in grasses, induction of physical defences is both specific to herbivore feeding, as opposed to mechanical damage, and may be dependant on the amount of damage imposed. Furthermore, we show that the magnitude of the induction response is sufficient to deter further damage and affect herbivore performance. We compared silica induction in two grass species in response to vertebrate and invertebrate damage, and to mechanical defoliation. Induction was assessed at two levels of damage over 16 months. Foliar silica content did not increase in response to mechanical defoliation, but damage by either voles or locusts resulted in increases in silica content of over 400%. This increase deterred feeding by both voles and locusts. Silica induction in grasses due to repeated damage events over a prolonged period suggests a possible role for silica defence in the cyclical population fluctuations observed in many grass-feeding herbivores.     

Competition between heather and grasses on Scottish moorlands: Interacting effects of nutrient enrichment and grazing regime

Abstract. Considerable losses and degradation of heathlands (in moorlands and lowlands) have been reported across Europe, with Calluna vulgaris (heather) being replaced by other species, often grasses. Increasing atmospheric nitrogen deposition and overgrazing have been suggested as the driving factors behind this change. This possibility was investigated in a study of the interacting effects of nutrient inputs and grazing on heather and three grass species (Nardus stricta, Deschampsia cespitosa and D. flexuosa) in the field, on a moorland in northeastern Scotland. In addition, the interacting effects of increasing nutrients and Calluna canopy height on N. stricta and D. cespitosa were studied using turves in an outdoor experimental area. In the field, fencing had a larger effect than fertilizer on the growth of all species, except for N. stricta, the species most unpalatable to herbivores. Fencing led to an increase in the height of the Calluna canopy, which may reduce light availability for the grasses. In the turf experiment, the height of the Calluna canopy affected the diameter of the grass tussocks and percentage of green matter (i.e. live leaf material), with plants under the more closed Calluna canopies being smaller. This study suggests that the slow‐growing, evergreen Calluna is a more effective competitor than the faster growing grasses when it has a tall, intact canopy, even at increased levels of nutrient supply. However, overgrazing promotes gap formation in the Calluna canopy, providing an opportunity for grasses to take advantage of increased nutrients. Thus the conservation of heather moorlands requires an understanding of the grazing level which allows Calluna to maintain sufficient canopy structure to outcompete grasses for light.     

Abiotic constraints on the competitive ability of exotic and native grasses in a Pacific Northwest prairie

In prairie ecosystems, abiotic constraints on competition can structure plant communities; however, the extent to which competition between native and exotic plant species is constrained by environmental factors is still debated. The objective of our study was to use paired field and greenhouse experiments to evaluate the competitive dynamics between two native (Danthonia californica and Deschampsia cespitosa) and two exotic (Schedonorus arundinaceus and Lolium multiflorum) grass species under varying nutrient and moisture conditions in an upland prairie in the Willamette Valley, Oregon. We hypothesized the two invasive, exotic grasses would be more competitive under high-nutrient, moderate-moisture conditions, resulting in the displacement of native grasses from these environments. In the field, the experimental reduction of competition resulted in shorter, wider plants, but only the annual grass, Lolium multiflorum, produced more aboveground biomass when competition was reduced. In the greenhouse, the two exotic grasses produced more total biomass than the two native grasses. Competitive hierarchies were influenced by nutrient and/or moisture treatments for the two exotic grasses, but not for the two native grasses. L. multiflorum dominated competitive interactions with all other grasses across treatments. In general, S. arundinaceus dominated when in competition with native grasses, and D. cespitosa produced the most biomass in monoculture or under interspecific competition with the other native grass, D. californica. D. californica, D. cespitosa, and S. arundinaceus all produced more biomass in high-moisture, high-nutrient environments, and D. cespitosa, L. multiflorum, and S. arundinaceus allocated more biomass belowground in the low nutrient treatment. Taken together, these experiments suggest the competitive superiority of the exotic grasses, especially L. multiflorum, but, contrary to our hypothesis, the native grasses were not preferentially excluded from nutrient-rich, moderately wet environments. Laurel Pfeifer-Meister and Esther M. Cole contributed equally to this work.     

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.     

Lack of intraspecific variation in resistance to defoliation in a grass that evolved under light grazing pressure

Grass species frequently show marked intraspecific variation inmorphology and tolerance to defoliation. Remarkably, most of this evidencecomesfrom grasslands with long evolutionary history of grazing. Here, we explore theintraspecific variation in grazing tolerance and morphometric traits associatedwith grazing avoidance of Paspalum dilatatum (Poir.), agrass from the Flooding Pampa (Argentina), where grazing is a novel disturbancein evolutionary time. We performed a clipping experiment in a greenhouse withtwo populations from sites with contrasting short term grazing regime:continuous grazing and 20 year-old grazing exclosure. The populations did notdiffer in their tolerance to clipping, and showed minor differences in the wayclipping affected plant height, a trait associated with grazing avoidance. Ourresults indicate that there are exceptions to the generalized findings of highlevels of intraspecific variation in grazing resistance among populations ofgrasses. These exceptions may be associated to evolutionary history of grazing.     

Nitrogen transfer between herbivores and their forage species

Herbivores may increase the productivity of forage plants; however, this depends on the return of nutrients from faeces to the forage plants. The aim of this study was to test if nitrogen (N) from faeces is available to forage plants and whether the return of nutrients differs between plant species using ¹⁵N natural abundance in faeces and plant tissue. To investigate the effect of grazing on N transfer, we carried out a grazing experiment in wet and mesic tundra on high Arctic Spitsbergen using barnacle geese (Branta leucopsis) as the model herbivore. N inputs (from faeces) increased with grazing pressure at both the wet and mesic sites, with the greatest N input from faeces at the wet site. The δ¹⁵N ratio in plant tissue from grazed plots was enriched in mosses and the dwarf shrub species, reflecting the δ¹⁵N signature of faeces-derived N, but no such pattern was observed in the dominant grasses. This study demonstrates that the δ¹⁵N signature of faeces and forage species is a useful tool to explore how grazing impacts on N acquisition. Our findings suggest that plant species which acquire their N close to the soil surface (e.g. mosses) access more of the N from faeces than species with deeper root systems (e.g. grasses) suggesting a transfer of N from the preferred forage species to the mosses and dwarf shrubs, which are less preferred by the geese. In conclusion, the moss layer appears to disrupt the nitrogen return from herbivores to their forage species.     

·荒漠草地植物多样性对草食动物采食的响应机制

草食动物采食对草地植物多样性和生态系统功能的影响机制是放牧生态学研究的核心问题。该研究以内蒙古锡林郭勒盟苏尼特右旗荒漠草原的长期放牧控制实验为平台, 从既有草地植物多样性和动物偏食性两个层面系统地研究了荒漠草地植物多样性对草食动物采食的响应机制。结果显示: 1)荒漠草地植物对草食动物采食呈现4种响应模式: 放牧“隐没种”、放牧“敏感种”、放牧“无感种”、“绝对优势种”; 2)在群落尺度上, 物种多样性指数随放牧强度增加而减少, 与不放牧小区相比, 重度放牧(HG)与适度放牧(MG)小区植物多样性均下降, 且这一规律同样适用于功能群多样性, 灌木半灌木这一功能群内物种多样性对放牧干扰较敏感; 3)在草地既有植物的基础上, 以不放牧小区为参考系, 草食动物对植物功能群偏食性的排序为: 一二年生草本(AB) >多年生杂类草(PF) >灌木半灌木(SS) >多年生禾草(PG), 且偏食性物种主要分布于AB和PF中; 4)植物多样性与动物偏食性基本呈显著负相关关系(p < 0.05)。     

The effect of herbivores on genotypic diversity in a clonal aquatic plant

In clonal plants, vegetative parts may outcompete seeds in the absence of disturbance, limiting the build‐up of genotypic diversity through repeated seedling recruitment (RSR). Herbivory may provide disturbance and trigger establishment of strong colonizers (seeds) at the expense of strong competitors (clonal propagules). In the clonal aquatic fennel pondweed Potamogeton pectinatus, two distinct herbivore guilds may modify the dynamics of propagation. In winter, Bewick's swans may deplete patches of tubers, promoting seedling establishment in spring. In summer, seed consumption by waterfowl can reduce the density of viable seeds but grazing may also reduce tuber production and hence facilitate seedling establishment. This study is among the first to experimentally test herbivore impact on plant genotypic diversity. We assess the separate and combined effects of both herbivore guilds on genotypic diversity and structure of fennel pondweed beds. Using microsatellites, we genotyped P. pectinatus from an exclosure experiment and assessed the contribution of herbivory, dispersal and sexual reproduction to the population genetic structure. Despite the predominance of clonal propagation in P. pectinatus, we found considerable genotypic diversity. Within the experimental blocks, kinship among genets decreased with geographic distance, clearly identifying a role for RSR in the maintenance of genotypic diversity within the fennel pondweed beds. However, over a period of five years, none of the herbivory treatments affected genotypic diversity. Hence, sexual reproduction on a local scale is important in this putatively clonal plant and possibly sufficient to ensure a relatively high genotypic diversity even in the absence of herbivores. Although we cannot preclude a role of herbivory in shaping genotypic diversity of a clonal plant, after five years of exclusion of the two investigated herbivore guilds no measurable effect on genotypic diversity was detected.     

Fine‐tuning of defences and counter‐defences in a specialised plant–herbivore system

1. The plant–herbivore arms race has been postulated to be a major driver for generating biological and biochemical diversity on Earth. Herbivore feeding is reduced by the production of chemical and physical barriers, but increases plant resistance against subsequent attack. Accordingly, specialisation is predicted to be an outcome of herbivores being able to circumvent plant‐induced defences. 2. Using a specialised plant–herbivore system, in which adult chrysomelid beetles (Chrysochus auratus) feed on leaves and larvae feed on roots of dogbane (Apocynum spp.), this study investigated whether root latex and cardenolides are effective against the soil‐dwelling larvae, and whether such defences could be circumvented by the herbivore. 3. Across two Apocynum species, C. auratus larvae were not affected by latex production or cardenolide amounts and diversity. By contrast, cardenolide apolarity was detrimental to larval growth. Yet larval feeding decreased average root cardenolide apolarity in A. cannabinum and larvae performed better on those plants. Finally, above‐ground induction rendered the plants more toxic by increasing root cardenolide apolarity and maintaining it, even during subsequent larval herbivory. 4. Therefore, the intimate relationship and interaction between Chrysochus and Apocynum are maintained by a delicate balance of herbivore manipulation and plant chemical induction.     

Experimental demonstration of the antiherbivore effects of silica in grasses: impacts on foliage digestibility and vole growth rates

The impact of plant-based factors on the population dynamics of mammalian herbivores has been the subject of much debate in ecology, but the role of antiherbivore defences in grasses has received relatively little attention. Silica has been proposed as the primary defence in grasses and is thought to lead to increased abrasiveness of foliage so deterring feeding, as well as reducing foliage digestibility and herbivore performance. However, at present there is little direct experimental evidence to support these ideas. In this study, we tested the effects of manipulating silica levels on the abrasiveness of grasses and on the feeding preference and growth performance of field voles, specialist grass-feeding herbivores. Elevated silica levels did increase the abrasiveness of grasses and deterred feeding by voles. We also demonstrated, for the first time, that silica reduced the growth rates of both juvenile and mature female voles by reducing the nitrogen they could absorb from the foliage. Furthermore, we found that vole feeding leads to increased levels of silica in leaves, suggesting a dynamic feedback between grasses and their herbivores. We propose that silica induction due to vole grazing reduces vole performance and hence could contribute to cyclic dynamics in vole populations.     

Leaf litter decomposition differs among genotypes in a local <Emphasis Type="Italic">Betula pendula</Emphasis> population

Ecosystem processes, such as plant litter decomposition, are known to be partly genetically determined, but the magnitude of genetic variation within local populations is still poorly known. We used micropropagated field-grown saplings of 19 Betula pendula genotypes, representing genetic variation in a natural birch population, to examine (1) whether genotype can explain variation in leaf litter decomposition within a local plant population, and (2) whether genotypic variation in litter decomposition is associated with genotypic variation in other plant attributes. We found that a local B. pendula population can have substantial genotypic variation in leaf litter mass loss at the early stages of the decomposition process and that this variation can be associated with genotypic variation in herbivore resistance and leaf concentrations of soluble proteins and total nitrogen (N). Our results are among the first to show that fundamental ecosystem processes can be significantly affected by truly intraspecific genetic variation of a plant species.     

An attempt to restore a central European species‐rich mountain grassland through grazing

Abstract. This paper describes the effects of re‐establishing seasonal cattle grazing by 0.7 animal.ha^‐1 on vegetation in a long‐term abandoned, and partly degraded, semi‐natural mountain pasture in the ?umava National Park, Czech Republic. There was very uneven grazing intensity inside the locality, and grazing preference changed during the season: cattle grazed most of the time in productive but species‐poor Deschampsia cespitosa swards, but changed to a species‐rich Violion caninae stand in the middle of the summer. A species‐rich Carex rostrata community was only grazed at the end of the season. Species‐poor swards dominated by Nardus stricta and Carex brizoides were mainly used as resting areas. Both grazing and excluding from grazing had a negative effect on species diversity of the Deschampsia cespitosa swards. The soil seed bank contained only few species that are characteristic of mountain grassland communities, and seed dispersal of the target species by cattle dung was also found to be very limited. Thus both grazing and exclusion from grazing are probably of limited value for the restoration of species‐rich grasslands from species‐poor Deschampsia cespitosa swards in this case.     

Ontogenetic changes in anti-herbivore defensive traits in leaves of four Mediterranean co-occurring <Emphasis Type="Italic">Quercus</Emphasis> species

Investment in anti-herbivore defence in tree species has been one of the priority research topics in plant terrestrial ecology during the last decades. However, despite considerable experimental effort, interspecific differences in the ontogenetic trends in the investment in defence are still a matter of debate, as to date experimental evidence is contradictory. In the present work, insect herbivory levels were measured in seedlings and mature trees of four co-occurring Mediterranean Quercus species with differing leaf life spans, as well as several leaf characteristics that can determine herbivore preference. The measured leaf traits included nitrogen (N), fibre (cellulose, hemicellulose and lignin), total phenolic contents, leaf mass per unit area (LMA) and leaf thickness. The leaves of seedlings had a lower LMA and leaf thickness and lower concentrations of N and cellulose, but higher concentrations of lignin and phenols than those of mature trees. However, the loss of leaf area tended to be more severe for seedlings than for mature trees, although the differences were only significant for deciduous species. This constitutes a confirmation of the strong effects of physical traits on herbivore preferences. The greater resource limitations for defensive mechanisms in seedlings with respect to mature trees would explain that at intraspecific level we do observe a compromise between chemical and physical defences. As a result, seedlings rely on chemical rather than on physical defences.     

Variation in multiple traits of vegetative and reproductive seagrass tissues influences plant–herbivore interactions

Plant–herbivore interactions have strong ecological and evolutionary consequences, but have been traditionally overlooked in marine higher plants. Despite recent advances in seagrass ecology that highlight the importance of herbivory, the mechanisms that regulate the feeding behaviour of seagrass consumers remain largely unknown. Herbivores have been shown to reduce the sexual reproductive success of seagrasses through direct consumption of inflorescences and seeds, but we know little about intraspecific variation in susceptibility to grazing of different seagrass tissues. We contrasted the relative palatability of reproductive and vegetative tissues of the temperate seagrass Posidonia oceanica in the field, and we assessed the feeding preferences among these tissues of the main consumers of the plant, the fish Sarpa salpa and the urchin Paracentrotus lividus. Moreover, we identified the plant traits that explained the observed feeding behaviour. We provide strong evidence for herbivore selectivity among seagrass tissues. In the field, 70–90% of inflorescences were damaged by herbivores compared to 3–60% of leaves of similar age. In feeding assays, the urchin P. lividus showed over a twofold preference for reproductive tissue at various stages of development. By contrast, we detected no feeding activity on either leaves or inflorescences from the fish S. salpa, which is known to migrate to deeper waters soon after flowering starts and during the period of fruit maturation. Despite being the preferred food of urchins, inflorescences were chemically defended, had higher levels of phenolics and lower nutrient and calorific content than leaves. We experimentally demonstrated that leaf structural defences are the primary factor in determining urchin feeding preferences. Removal of plant structure results in a drastic shift in urchin selectivity towards the most nutritious and less chemically defended leaf tissue, indicating that multiple mechanisms of defence to herbivory may coexist in seagrasses.     

Palatability factors and nutritive values of the food of buffaloes (Syncerus caffer) in Uganda

The food preferences of tame buffaloes grazing different vegetation types were recorded. The effects of fire and rainfall were investigated. By initiating a new growth cycle, fire altered the food habits of large herbivores and also the distribution of small mammals. Seasonal variations in the effect of rainfall on plant growth explains why preferences did not change in proportion to precipitation. The more important grasses were collected and their parts analysed physically and chemically. Their stem to leaf ratio is considered to have influenced the species eaten by coarse grazers such as buffalo. A broad correlation was obtained between buffalo preferences and the percentage green leaf of Hyparrhenia filipendula and Themeda triandra. This correlation was improved when preferences were compared with the number of grams green leaf carbohydrate per 100 g dry grass. There was a negative response to the silica present in dead leaves, but stem fibre appeared to have little effect on preferences. A grazing animal experiences a combination of attractants and repellants, i.e. green leaf carbohydrate and protein less stem fibre and dead leaf silica. Other palatability factors were noted, in particular tannins and oils in grasses of the tribe Andropogoneae. These may act as repellants, possibly only when the grazing animal has alternative vegetation from which to choose. The mineral content of grasses appeared to be adequate and calcium and phosphorus were present in about the required proportions. The observations support the belief that selective grazing enables a herbivore to consume a diet of significantly higher nutritive value than that of the average sward. Browsing and new growth after fire help compensate for the low nutritive value of pasture during the dry season. The nutritive value of the diet is considered to be the proximal factor in the seasonal cycle of reproduction of the buffalo and probably the ultimate factor in the reproduction of certain other ungulates and rodents.     

Specificity in Mesograzer-Induced Defences in Seagrasses

Grazing-induced plant defences that reduce palatability to herbivores are widespread in terrestrial plants and seaweeds, but they have not yet been reported in seagrasses. We investigated the ability of two seagrass species to induce defences in response to direct grazing by three associated mesograzers. Specifically, we conducted feeding-assayed induction experiments to examine how mesograzer-specific grazing impact affects seagrass induction of defences within the context of the optimal defence theory. We found that the amphipod Gammarus insensibilis and the isopod Idotea chelipes exerted a low-intensity grazing on older blades of the seagrass Cymodocea nodosa, which reflects a weak grazing impact that may explain the lack of inducible defences. The isopod Synischia hectica exerted the strongest grazing impact on C. nodosa via high-intensity feeding on young blades with a higher fitness value. This isopod grazing induced defences in C. nodosa as indicated by a consistently lower consumption of blades previously grazed for 5, 12 and 16 days. The lower consumption was maintained when offered tissues with no plant structure (agar-reconstituted food), but showing a reduced size of the previous grazing effect. This indicates that structural traits act in combination with chemical traits to reduce seagrass palatability to the isopod. Increase in total phenolics but not in C:N ratio and total nitrogen of grazed C. nodosa suggests chemical defences rather than a modified nutritional quality as primarily induced chemical traits. We detected no induction of defences in Zostera noltei, which showed the ability to replace moderate losses of young biomass to mesograzers via compensatory growth. Our study provides the first experimental evidence of induction of defences against meso-herbivory that reduce further consumption in seagrasses. It also emphasizes the relevance of grazer identity in determining the level of grazing impact triggering resistance and compensatory responses of different seagrass species.     

Restoration of grazing management and its effect on vegetation in an upland grassland

Question: How are plant species and functional group composition, and potential sward height affected by implementation of different grazing regimes on previously abandoned semi-natural grassland? Location: The Jizerské mountains, northern Czech Republic. Methods: We established a randomized block experiment with the following treatments: unmanaged control (U), intensive (IG) and extensive (EG) continuous grazing, first cut followed by intensive (ICG) and first cut followed by extensive (ECG) continuous grazing for the rest of the growing season. The percentage cover of all vascular plant species was recorded in 40 permanent plots. Results: Total plant species richness increased in all managed treatments, whereas species number was reduced in U at the end of the experiment. Tall forbs (Aegopodium podagraria, Galium album, Anthriscus sylvestris, Cirsium arvense) as well as tall grasses (Elytrigia repens and Alopecurus pratensis) were more abundant in U. Species associated with both grazing treatments (IG, EG) were Dactylis glomerata, Festuca rubra agg. and Phleum pratense. Agrostis capillaris, Taraxacum spp., Trifolium repens, Ranunculus repens and Cirsium vulgare were promoted by ECG and ICG. Abundance of tall grasses and tall forbs reflected the intensity of management in the order U>EG, ECG>IG, ICG. Prostrate forbs, on the other hand, increased their cover with increasing intensity: ICG>IG>ECG>EG. Conclusions: Plant species composition of semi-natural grasslands is affected by the defoliation regime. Continuous grazing on abandoned grassland alters the sward structure towards a permanent pasture with short, light-sensitive grasses and prostrate forbs. To maintain or enhance plant species richness in semi-natural grasslands, understanding the effects of different grazing regimes on plant species composition is necessary.     

Fungal endophytes of native grasses decrease insect herbivore preference and performance

Endophytic fungal symbionts of grasses are well known for their protective benefit of herbivory reduction. However, the majority of studies on endophyte–grass symbioses have been conducted on economically important, agricultural species—particularly tall fescue (Lolium arundinaceum) and perennial ryegrass (Lolium perenne)—raising the hypothesis that strong benefits are the product of artificial selection. We examined whether fungal endophytes found in natural populations of native grass species deterred insect herbivores. By testing several native grass–endophyte symbiota, we examined phylogenetic signals in the effects of endophytes on insects and compared the relative importance of herbivore and symbiotum identity in the outcome of the interactions. Preference was assessed using three herbivore species [Spodoptera frugiperda (Lepidoptera), Schistocerca americana (Orthoptera), Rhopalosiphum padi (Hemiptera)] and ten native symbiota, which spanned seven grass genera. We also assessed herbivore performance in a no choice experiment for five native symbiota against S. frugiperda. We compared greenhouse and laboratory trials with natural levels of herbivory measured in experimental field populations. In all cases, we included the agronomic grass species, L. arundinaceum, to compare with results from the native grasses. Both in the field and in experimental trials, herbivores showed a significant preference for endophyte-free plant material for the majority of native grasses, with up to three times lower herbivory for endophyte-symbiotic plants; however, the degree of response depended on the identity of the herbivore species. Endophyte presence also significantly reduced performance of S. frugiperda for the majority of grass species. In contrast, the endophyte in L. arundinaceum had few significant anti-herbivore effects, except for a reduction in herbivory at one of two field sites. Our results demonstrate that the mechanisms by which native symbionts deter herbivores are at least as potent as those in model agricultural systems, despite the absence of artificial selection.     

高寒草地植物群落关键种对不同放牧家畜组合放牧的响应

如何通过合理的利用方式提高高寒草地管理水平,实现其可持续利用一直是草地生态学领域的研究热点。为明确不同放牧家畜组合下高寒草地植物群落关键种的演替规律及其驱动因素,基于中等放牧强度设置了不同放牧家畜组合放牧样地（牦牛单牧、藏羊单牧、牦牛藏羊1 ： 2混牧、牦牛藏羊1 ： 4混牧、牦牛藏羊1 ： 6混牧）以及围封样地,并在连续放牧处理7年后系统分析了不同放牧家畜组合下植物群落特征与土壤理化性质变化,以期确定最优放牧组合。结果表明：（1）不同放牧家畜组合均会显著降低高寒草地植物盖度,但对其物种丰富度与多样性指数的影响并不显著。（2）牦牛藏羊1 ： 2混牧下植物群落特征与禁牧处理下植物群落特征较为相似,且牦牛藏羊1 ： 2混牧下的植物群落稳定性最强、组织水平最高。（3）牦牛单牧与1 ： 2混牧下关键种为矮生嵩草;藏羊单牧下关键种为天山针茅;1 ： 4混牧下关键种为星毛委陵菜;1 ： 6混牧与围封下关键种为赖草。（4）围封能够显著降低土壤容重,藏羊单牧则会显著增加土壤容重与土壤速效氮、磷含量。（5）土壤速效氮、容重与含水量是驱动不同放牧家畜组合下植物群落关键种演替的重要理化因子。综上所述,中等放牧强度下,牦牛藏羊1 ： 2混牧是青藏高原高寒草地较为理想的利用方式。此外,相较于单纯增加草地物种丰富度或多样性,建立植物群落物种之间的有效关联是提高青藏高原高寒草地管理水平的另一关键途径。