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Endophytic fungi, especially asexual, systemic endophytes in grasses, are generally viewed as plant mutualists, mainly through the action of mycotoxins, such as alkaloids in infected grasses, which protect the host plant from herbivores. Most of the evidence for the defensive mutualism concept is derived from studies of agronomic grass cultivars, which may be atypical of many endophyte-host interactions. I argue that endophytes in native plants, even asexual, seed-borne ones, rarely act as defensive mutualists. In contrast to domesticated grasses where infection frequencies of highly toxic plants often approach 100%, natural grass populations are usually mosaics of uninfected and infected plants. The latter, however, usually vary enormously in alkaloid levels, from none to levels that may affect herbivores. This variation may result from diverse endophyte and host genotypic combinations that are maintained by changing selective pressures, such as competition, herbivory and abiotic factors. Other processes, such as spatial structuring of host populations and endophytes that act as reproductive parasites of their hosts, may maintain infection levels of seed-borne endophytes in natural populations, without the endophyte acting as a mutualist.

A characteristic feature of fish is the wide physiological range of blood parameters and also the large individual variations. The aim of this study was to compare the haematological profile, glucose and lactate levels of four teleost fish species (Gobius niger, Mugil cephalus, Sparus aurata, Dicentrarchus labrax) and to establish the similarities and differences between these species which are widely present in the Tyrrhenian Sea. To this end, glucose, lactate and complete haematological profiles were determined for 25 fish from each species. Statistical analysis confirmed statistical differences in blood parameters among the four species. Our findings show a lower level of glucose and higher levels of lactate, red blood cells and haemoglobin in M. cephalus with respect to the other species. White blood cell and thrombocyte counts have the same trend and result higher in S. aurata. The differences found in this study can be attributed to the feeding behaviour, life style and adaptation of the different fish species to the habitat in which they dwell.

Herbivore attack is widely known to reduce food quality and to increase chemical defenses and other traits responsible for herbivore resistance. Inducible defenses are commonly thought to allow plants to forgo the costs of defense when not needed; however, neither their defensive function (increasing a plant's fitness) nor their cost-savings function have been demonstrated in nature. The root-produced toxin nicotine increases after herbivore attack in the native, postfire annual Nicotiana attenuata and is internally activated by the wound hormone, jasmonic acid. I treated the roots of plants with the methyl ester of this hormone (MeJA) to elicit a response in one member of each of 745 matched pairs of plants growing in native populations with different probabilities of attack from herbivores, and measured the lifetime production of viable seed. In populations with intermediate rates of attack, induced plants were attacked less often by herbivores and survived to produce more seed than did their uninduced counterparts. Previous induction did not significantly increase the fitness of plants suffering high rates of attack. However, if plants had not been attacked, induced plants produced less seed than did their uninduced counterparts. Jasmonate-induced responses function as defenses but are costly, and inducibility allows this species to forgo these costs when the defenses are unnecessary

Plant resistance to herbivores was induced in a field experiment to evaluate the consequences of induced responses for subsequent herbivory and plant fitness. Induction early in the season resulted in halving of herbivory by chewing herbivores and a reduction in the abundance of phloem-feeding aphids when compared with controls. A correlate of lifetime plant fitness, seed mass, was enhanced by over 60 percent for individuals that were induced

Herbivores can often control plant dynamics by mediating positive feedbacks in plant species' influence on nutrient cycling. In a 7-yr field experiment in a nitrogen-limited Minnesota oak savanna, we tested whether herbivores accelerated or decelerated nitrogen (N) cycling through their effects on plants. We measured effects of excluding insect (primarily Orthoptera and Homoptera) and mammalian herbivores (primarily white-tailed deer, Odocoileus virginianus) on above- and belowground plant biomass, plant species composition, plant tissue nitrogen concentration, available soil N and light, and total N and carbon (C) in different pools (soil, roots, litter, etc.). Herbivore exclusion strongly increased cover and biomass of the legume Lathyrus venosus and a few species of woody plants. These effects were associated with greater aboveground standing crop, reduced belowground standing crop, and reduced light penetration to the ground surface. Herbivore exclusion also modified N cycling through greater N content of live, aboveground plant tissue early in the growing season and of litter and belowground tissue late in the growing season. Herbivore exclusion also increased soil nitrate and total available N concentrations but did not alter total soil or plant N. Our results support the hypothesis that herbivores indirectly decelerate N cycling by decreasing the abundance of plant species with nitrogen-rich tissues. However, other factors, such as disturbance from fire, may mediate herbivore effects on long-term changes in N and C pools. Herbivores may therefore indirectly control productivity, N cycling, and succession by consuming nitrogen-fixing and woody plants that have strong effects on plant resources (e.g., nitrogen and light).

Empirical research has not supported the prediction that populations of terrestrial herbivorous arthropods are regulated solely by their natural enemies. Instead, both natural enemies (top-down effects) and resources (bottom-up effects) may play important regulatory roles. This review evaluates the hypothesis that higher-order predators may constrain the top-down control of herbivore populations. Natural enemies of herbivorous arthropods generally are not top predators within terrestrial food webs. Insect pathogens and entomopathogenic nematodes inhabiting the soil may be attacked by diverse micro- and mesofauna. Predatory and parasitic insects are attacked by their own suite of predators, parasitoids, and pathogens. The view of natural enemy ecology that has emerged from laboratory studies, where natural enemies are often isolated from all elements of the biotic community except for their hosts or prey, may be an unreliable guide to field dynamics. Experimental work suggests that interactions of biological control agents with their own natural enemies can disrupt the effective control of herbivore populations. Disruption has been observed experimentally in interactions of bacteria with bacteriophages, nematodes with nematophagous fungi, parasitoids with predators, parasitoids with hyperparasitoids, and predators with other predators. Higher-order predators have been little studied; manipulative field experiments will be especially valuable in furthering our understanding of their roles in arthropod communities.

In response to insect feeding on the leaves, cotton (Gossypium hirsutum L.) plants release elevated levels of volatiles, which can serve as a chemical signal that attracts natural enemies of the herbivore to the damaged plant. Pulse-labeling experiments with [13C]CO2 demonstrated that many of the volatiles released, including the acyclic terpenes (E,E)-alpha-farnesene, (E)-beta-farnesene, (E)-beta-ocimene, linalool,(E)-4,8-dimethyl-1,3,7-nonatriene, and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetrane, as well as the shikimate pathway product indole, are biosynthesized de novo following insect damage. However, other volatile constituents, including several cyclic terpenes, butyrates, and green leaf volatiles of the lipoxygenase pathway are released from storage or synthesized from stored intermediates. Analysis of volatiles from artificially damaged plants, with and without beet armyworm (Spodoptera exigua Hubner) oral secretions exogenously applied to the leaves, as well as volatiles from beet armyworm-damaged and -undamaged control plants, demonstrated that the application of caterpillar oral secretions increased both the production and release of several volatiles that are synthesized de novo in response to insect feeding. These results establish that the plant plays an active and dynamic role in mediating the interaction between herbivores and natural enemies of herbivores

Within large grassland ecosystems, climatic and topographic gradients are considered the primary controls of soil processes. Ungulates also can influence soil dynamics; however the relative contribution of large herbivores to controlling grassland soil processes remains largely unknown. In this study, we compared the effects of native migratory ungulates and variable site (\"landscape\") conditions, caused by combined climatic and topographic variability, on grassland of the northern winter range of Yellowstone National Park by determining soil C and N dynamics inside and outside 33-37 yr exclosures at seven diverse sites. Sites included hilltop, slope, and slope bottom positions across a climatic gradient and represented among the driest and wettest grasslands on the northern winter range. We performed two experiments: (1) a 12-mo in situ net N mineralization study and (2) a long-term (62-wk) laboratory incubation to measure potential N mineralization and microbial respiration. Results from the in situ experiment indicated that average net N mineralization among grazed plots (3.8 g N· m-2· yr-1) was double that of fenced, ungrazed plots (1.9 g N· m-2· yr-1). Mean grazer enhancement of net N mineralization across sites (1.9 g N· m-2· yr-1) approached the maximum difference in net N mineralization among fenced plots (2.2 g N· m-2· yr-1), i.e., the greatest landscape effect observed. Furthermore, ungulates substantially increased between-site variation in mineralization; grazed grassland, 1 SD = 2.2 g N· m-2· yr-1, fenced grassland, 1 SD = 2.2 g N· m-2· yr-1. In the long-term incubation, potential microbial respiration and net N mineralization were positively related to total soil C and N content, respectively. There was greater variation in potential respiration and net N mineralization early in the incubation, when labile material was processed, compared to late in the incubation, when more recalcitrant substrate was processed, suggesting that between-site variation in labile organic matter was greater than that of recalcitrant material. Herbivores improved the organic matter quality of soil, increasing the labile fractions and reducing the recalcitrant fractions. Grazers reduced C respired/N mineralized ratios, an index of microbial N immobilization, by an average of 21%. However, the largest landscape influence on the immobilization index was 13-fold greater than the grazer effect. Given that the greatest landscape influence on in situ net mineralization (2.2 g N· m-2· yr-1) was similar to the average grazer impact on that rate (1.9 g N· m-2· yr-1), we hypothesize that the landscape effect on field N availability was primarily caused by variation in microbial immobilization, while the grazing effect was primarily due to stimulation of gross mineralization. These results indicate that the relative importance of ungulates in controlling soil N cycling may be more important than previously suspected for grasslands supporting large herds of migratory ungulates, and that the dominant mechanisms underlying the landscape and ungulate influences on soil mineral fluxes may differ.

Predators of herbivorous animals can affect plant populations by altering herbivore density, behavior, or both. To test whether the indirect effect of predators on plants arises from density or behavioral responses in a herbivore population, we experimentally examined the dynamics of terrestrial food chains comprised of old field plants, leaf-chewing grasshoppers, and spider predators in Northeast Connecticut. To separate the effects of predators on herbivore density from the effects on herbivore behavior, we created two classes of spiders: risk spiders that had their feeding mouth parts glued to render them incapable of killing prey and predator spiders that remained unmanipulated. We found that the effect of predators on plants resulted from predator-induced changes in herbivore behavior (shifts in activity time and diet selection) rather than from predator-induced changes in grasshopper density. Neither predator nor risk spiders had a significant effect on grasshopper density relative to a control. This demonstrates that the behavioral response of prey to predators can have a strong impact on the dynamics of terrestrial food chains. The results make a compelling case to examine behavioral as well as density effects in theoretical and empirical research on food chain dynamics

Two potential outcomes of a coevolutionary interaction are an escalating arms race and stable cycling. The general expectation has been that arms races predominate in cases of polygenic inheritance of resistance traits and permanent cycling predominates in cases in which resistance is controlled by major genes. In the interaction between Depressaria pastinacella, the parsnip webworm, and Pastinaca sativa, the wild parsnip, traits for plant resistance to insect herbivory (production of defensive furanocoumarins) as well as traits for herbivore \"virulence\" (ability to metabolize furanocoumarins) are characterized by continuous heritable variation. Furanocoumarin production in plants and rates of metabolism in insects were compared among four midwestern populations; these traits then were classified into four clusters describing multitrait phenotypes occurring in all or most of the populations. When the frequency of plant phenotypes belonging to each of the clusters is compared with the frequency of the insect phenotypes in each of the clusters across populations, a remarkable degree of frequency matching is revealed in three of the populations. That frequencies of phenotypes vary among populations is consistent with the fact that spatial variation occurs in the temporal cycling of phenotypes; such processes contribute in generating a geographic mosaic in this coevolutionary interaction on the landscape scale. Comparisons of contemporary plant phenotype distributions with phenotypes of herbarium specimens collected 9-125 years ago from across a similar latitudinal gradient, however, suggest that for at least one resistance trait-sphondin concentration-interactions with webworms have led to escalatory change