Mutualism

Here, the coevolution of mycorrhizal fungi and roots is assessed in the light of evidence now available, from palaeobotanical and morphological studies and the analysis of DNA-based phylogenies. The first bryophyte-like land plants, in the early Devonian (400 million years ago), had endophytic associations resembling vesiculararbuscular mycorrhizas (VAM) even before roots evolved. Mycorrhizal evolution would have progressed from endophytic hyphae towards balanced associations where partners were interdependent due to the exchange of limiting energy and nutrient resources. Most mycorrhizas are mutualistic, but in some cases the trend for increasing plant control of fungi culminates in the exploitative mycorrhizas of achlorophyllous, mycoheterotrophic plants. Ectomycorrhizal, ericoid and orchid mycorrhizas, as well as nonmycorrhizal roots, evolved during the period of rapid angiosperm radiation in the Cretaceous. It is hypothesised that roots gradually evolved from rhizomes to provide more suitable habitats for mycorrhizal fungi and provide plants with complex branching and leaves with water and nutrients. Selection pressures have caused the morphological divergence of roots with different types of mycorrizas. Root cortex thickness and exodermis suberization are greatest in obligately mycorrhizal plants, while nonmycorrhizal plants tend to have fine roots, with more roots hairs and relatively advanced chemical defences. Major coevolutionary trends and the relative success of plants with different root types are discussed.

The main theories of biodiversity either neglect species interactions 1,2 or assume that species interact randomly with each other 3,4 . However, recent empirical work has revealed that ecological networks are highly structured , and the lack of a theory that takes into account the structure of interactions precludes further assessment of the implications of such network patterns for biodiversity. Here we use a combination of analytical and empirical approaches to quantify the influence of network architecture on the number of coexisting species. As a case study we consider mutualistic networks between plants and their animal pollinators or seed dispersers 5,8-11 . These networks have been found to be highly nested 5 , with the more specialist species interacting only with proper subsets of the species that interact with the more generalist. We show that nestedness reduces effective interspecific competition and enhances the number of coexisting species. Furthermore, we show that a nested network will naturally emerge if new species are more likely to enter the community where they have minimal competitive load. Nested networks seem to occur in many biological and social contexts 12-14 , suggesting that our results are relevant in a wide range of fields.

The moment the soil enters into contact with an earthworm, both superficially and intemally, physicochemical and biological changes take place. The drilosphere represents the whole soil volume under earthworm influence. Thus it includes the body surfaces, the gut and all the internal features of the worm that are in contact with the ingested soil, as well as the external structures (casts, burrows, middens) created by earthworm activities. The extent of the drilosphere and its particular characteristics depend on the species and ecological categories of the earthworm community present as well as the spatial and temporal scale of interest. Spatially, the drilosphere can interact with other soil functional domains and lead to significant changes in the litter system or detritusphere (generally decreasing litter stocks) and the rhizosphere (affecting both root biomass and density), the two main sources of organic matter (OM) additions to the soil, as well as in the aggregatusphere and the porosphere. Drilosphere effects on microbial activity and OM decomposition can be completely different (and opposite) depending on the spatio-temporal scale of observation. At the level of the gut, microbial activity is dramatically stimulated in a matter of a few hours via a mutualistic digestion system. In this process, water and soluble-C in the form of intestinal mucus (the Kiss) produced by the earthworm (Prince Charming) awakens the dormant microflora (Sleeping Beauties), thereby increasing decomposition of the stable forms of soil OM ingested. During gut passage populations of other organisms (e.g. protozoa, nematodes, fungi) may decline with digestion, although these organisms probably form a minor component of the earthworm's energy needs. In the casts and on the burrow walls, the abundant nutrient resources for soil microflora continue the priming effect of the gut, increasing over a short time period mineralization rates and plant nutrient bio-availability. However as castings, particularly of the 'compacting group', and burrow walls begin to dry and stabilize with age (days to weeks), OM decomposition, nutrient mineralization and microbial activity decrease, often reaching levels lower than uningested soil due to 'protection'. Finally at the scale of years to decades and soil profile, it,appears that the drilosphere can exert an important regulation on OM incorporation and tumover rates, and soil C stocks. O 2000 Editions scientifiques et médicales Elsevier SAS I ~~~~~~~~~~

Herbivorous and carnivorous arthropods use plant volatiles when foraging for food. In response to herbivory, plants emit a blend that may be quantitatively and qualitatively different from the blend emitted when intact. This induced volatile blend alters the interactions of the plant with its environment. We review recent developments regarding the induction mechanism as well as the ecological consequences in a multitrophic and evolutionary context. It has been well established that carnivores (predators and parasitoids) are attracted by the volatiles induced by their herbivorous victims. This concerns an active plant response. In the case of attraction of predators, this is likely to result in a fitness benefit to the plant, because through consumption a predator removes the herbivores from the plant. However, the benefit to the plant is less clear when parasitoids are attracted, because parasitisation does usually not result in an instantaneous or in a complete termination of consumption by the herbivore. Recently, empirical evidence has been obtained that shows that the plant's response can increase plant fitness, in terms of seed production, due to a reduced consumption rate of parasitized herbivores. However, apart from a benefit from attracting carnivores, the induced volatiles can have a serious cost because there is an increasing number of studies that show that herbivores can be attracted. However, this does not necessarily result in settlement of the herbivores on the emitting plant. The presence of cues from herbivores and/or carnivores that indicate that the plant is a competitor-and/or enemy-dense space, may lead to an avoidance response. Thus, the benefit of emission of induced volatiles is likely to depend on the prevailing faunal composition. Whether plants can adjust their response and influence the emission of the induced volatiles, taking the prevalent environmental conditions into account, is an interesting question that needs to be addressed. The induced volatiles may also affect interactions of the emitting plant with its neighbours, e.g., through altered competitive ability or by the neighbour exploiting the emitted information.

Dark septate endophytes (DSE) are a miscellaneous group of ascomycetous anamorphic fungi that colonize root tissues intracellularly and intercellularly. The limited selection of studies quoted here exemplifies the range of host responses to symbiotic DSE fungi. Like mycorrhizal associations, DSE associations vary from negative to neutral and positive when measured by host performance or host tissue nutrient concentrations. This range of host responses is partially attributable to variation between different fungus taxa and strains. Similarly, hosts differ in their responses to a single DSE strain. Experimental conditions may also govern the nature of the symbiotic association. It is concluded that DSE are capable of forming mutualistic associations functionally similar to mycorrhizas. If the variation in host response to mycorrhizal fungi is considered to represent a continuum ranging from parasitism to mutualism, DSE symbiosis must be considered mycorrhizal, at least under some conditions.

Ecologists and evolutionary biologists are broadly interested in how the interactions among organisms influence their abundance, distribution, phenotypes, and genotypic composition. Recently, we have seen a growing appreciation of how multispecies interactions can act synergistically or antagonistically to alter the ecological and evolutionary outcomes of interactions in ways that differ fundamentally from outcomes predicted by pairwise interactions. Here, we review the evidence for criteria identified to detect community-based, diffuse coevolution. These criteria include (a) the presence of genetic correlations between traits involved in multiple interactions, (b) interactions with one species that alter the likelihood or intensity of interactions with other species, and (c) nonadditive combined effects of multiple interactors. In addition, we review the evidence that multispecies interactions have demographic consequences for populations, as well as evolutionary consequences. Finally, we explore the experimental and analytical techniques, and their limitations, used in the study of multispecies interactions. Throughout, we discuss areas in particular need of future research.

1. Almost all plants are engaged in symbiotic relationships with mycorrhizal fungi. These soil fungi can promote plant growth by supplying limiting nutrients to plant roots in return for plant assimilates. 2. Many mycorrhizal fungi are not host specific and one fungal individual can colonize and interconnect a considerable number of plants. The existence of these so-called mycorrhizal networks implies that fungi have the potential to facilitate growth of other plants and distribute resources among plants irrespective of their size, status or identity. In this paper, we explore the significance of mycorrhizal fungal networks for individual plants and for plant communities. 3. We address the following questions: (i) are all plant species benefitting from mycorrhizal networks, (ii) is benefit dependent on the size or age of a plant, (iii) is fungal support related to the relative dominance of plants in a community, (iv) are there host dependent barriers and physiological constraints for support and (v) what is the impact of mycorrhizal networks on plant-plant interactions and plant community dynamics? Moreover, using a review of published studies, we test whether mycorrhizal networks facilitate growth of small seedlings that establish between or near larger plants. 4. We found 60 cases where seedling species were grown together with larger plants with or without mycorrhizal fungal networks. Mycorrhizal networks promoted seedling growth in 48% of the cases (for 21 seedling species), while negative effects (25%) and no effects (27%) were also common. Seedlings associating with ectomycorrhizal fungi benefitted in the majority of the cases while effects on seedlings associating with arbuscular mycorrhizal fungi were more variable. Thus, the facilitative effects of mycorrhizal fungal networks depend on seedling species identity, mycorrhizal identity, plant species combinations and study system. We present a number of hypothetical scenarios that can explain the results based on cost-benefit relationship of individual members in a network. 5. Synthesis. Overall, this review shows that mycorrhizal networks play a key role in plant communities by facilitating and influencing seedling establishment, by altering plant-plant interactions and by supplying and recycling nutrients.

Most mycorrhizas are ' balanced ' mutualistic associations in which the fungus and plant exchange commodities required for their growth and survival. Myco-heterotrophic plants have 'exploitative ' mycorrhizas where transfer processes apparently benefit only plants. Exploitative associations are symbiotic (in the broad sense), but are not mutualistic. A new definition of mycorrhizas that encompasses all types of these associations while excluding other plant-fungus interactions is provided. This definition recognises the importance of nutrient transfer at an interface resulting from synchronised plant-fungus development. The diversity of interactions between mycorrhizal fungi and plants is considered. Mycorrhizal fungi also function as endophytes, necrotrophs and antagonists of host or non-host plants, with roles that vary during the lifespan of their associations. It is recommended that mycorrhizal associations are defined and classified primarily by anatomical criteria regulated by the host plant. A revised classification scheme for types and categories of mycorrhizal associations defined by these criteria is proposed. The main categories of vesicular-arbuscular mycorrhizal associations (VAM) are 'linear' or 'coiling ', and of ectomycorrhizal associations (ECM) are ' epidermal ' or 'cortical '. Subcategories of coiling VAM and epidermal ECM occur in certain host plants. Fungus-controlled features result in ' morphotypes ' within categories of VAM and ECM. Arbutoid and monotropoid associations should be considered subcategories of epidermal ECM and ectendomycorrhizas should be relegated to an ECM morphotype. Both arbuscules and vesicles define mycorrhizas formed by glomeromycotan fungi. A new classification scheme for categories, subcategories and morphotypes of mycorrhizal associations is provided.

Plant-animal mutualisms are characterized by weak or asymmetric mutual dependences between interacting species, a feature that could increase community stability. If invasive species integrate into mutualistic webs, they may alter web structure, with consequences for species persistence. However, the effect of alien mutualists on the architecture of plant-pollinator webs remains largely unexplored. We analyzed the extent of mutual dependency between interacting species, as a measure of mutualism strength, and the connectivity of 10 paired plant-pollinator webs, eight from forests of the southern Andes and two from oceanic islands, with different incidences of alien species. Highly invaded webs exhibited weaker mutualism than less-invaded webs. This potential increase in network stability was the result of a disproportionate increase in the importance and participation of alien species in the most asymmetric interactions. The integration of alien mutualists did not alter overall network connectivity, but links were transferred from generalist native species to super-generalist alien species during invasion. Therefore, connectivity among native species declined in highly invaded webs. These modifications in the structure of pollination webs, due to dominance of alien mutualists, can leave many native species subject to novel ecological and evolutionary dynamics.

Plants are rife with bacteria and fungi that colonize roots and shoots both externally and internally. By providing novel nutritional and defense pathways and influencing plant biochemical pathways, microbes can fundamentally alter plant phenotypes. Here we review the widespread nature of microbially mediated plant functional traits. We highlight that there is likely fitness conflict between hosts and symbionts and that fitness outcomes can depend on partner genotypes and ecological factors. Microbes may influence ecosystems through their effects on the functional trait values and population dynamics of their plant hosts. These effects may feed back on symbiont evolution by altering transmission rates of symbionts and scale up to ecosystem processes and services. We end by proposing new avenues of research in this emerging field. 23 Annu. Rev. Ecol. Evol. Syst. 2011.42:23-46. Downloaded from www.annualreviews.org by 76.171.15.208 on 09/27/12. For personal use only.

The effects of species declines and extinction on biotic interactions remain poorly understood. The loss of a species is expected to result in the loss of other species that depend on it (coextinction), leading to cascading effects across trophic levels. Such effects are likely to be most severe in mutualistic and parasitic interactions. Indeed, models suggest that coextinction may be the most common form of biodiversity loss. Paradoxically, few historical or contemporary coextinction events have actually been recorded. We review the current knowledge of coextinction by: (i) considering plausible explanations for the discrepancy between predicted and observed coextinction rates; (ii) exploring the potential consequences of coextinctions; (iii) discussing the interactions and synergies between coextinction and other drivers of species loss, particularly climate change; and (iv) suggesting the way forward for understanding the phenomenon of coextinction, which may well be the most ins...

Previous studies have shown that arbuscular mycorrhizal fungi (AMF) can influence plant diversity and ecosystem productivity. However, little is known about the effects of AMF and different AMF taxa on other important community properties such as nutrient acquisition, plant survival and soil structure. • We established experimental grassland microcosms and tested the impact of AMF and of different AMF taxa on a number of grassland characteristics. We also tested whether plant species benefited from the same or different AMF taxa in subsequent growing seasons. • AMF enhanced phosphorus acquisition, soil aggregation and survival of several plant species, but AMF did not increase total plant productivity. Moreover, AMF increased nitrogen acquisition by some plant species, but AMF had no effect on total N uptake by the plant community. Plant growth responses to AMF were temporally variable and some plant species obtained the highest biomass with different AMF in different years. Hence the results indicate that it may be beneficial for a plant to be colonized by different AMF taxa in different seasons. • This study shows that AMF play a key role in grassland by improving plant nutrition and soil structure, and by regulating the make-up of the plant community.

We isolated and examined two endophytic fungi for their potential to secrete phytohormones viz. gibberellins (GAs) and indoleacetic acid (IAA) and mitigate abiotic stresses like salinity and drought. The endophytic fungi Phoma glomerata LWL2 and Penicillium sp. LWL3 significantly promoted the shoot and allied growth attributes of GAs-deficient dwarf mutant Waito-C and Dongjin-beyo rice. Analysis of the pure cultures of these endophytic fungi showed biologically active GAs (GA 1 , GA 3 , GA 4 and GA 7 ) in various quantities. The cultures of P. glomerata and Penicillium sp. also contained IAA. The culture application and endophytic-association with host-cucumber plants significantly increased the plant biomass and related growth parameters under sodium chloride and polyethylene glycol induced salinity and drought stress as compared to control plants. The endophytic symbiosis resulted in significantly higher assimilation of essential nutrients like potassium, calcium and magnesium as compared to control plants during salinity stress. Endophytic-association reduced the sodium toxicity and promoted the host-benefit ratio in

A comprehensive and critical review of all available literature on associations between Australian lycaenid butterflies and ants was undertaken to establish an accurate database of the partners involved. Collections and observations of lycaenids and ants were used to augment this review, resulting in a significant number of newly documented association (and non-association) records. Twenty published records considered to be erroneous or doubtful are noted, with justifications given for their deletion from the association database. In total, 265 different associations between lycaenids and ants, plus 65 non-attendance records are documented for Australia. Nearly 80% of the lycaenid species in Australia, for which the early stages are known, are recorded associating with ants and half of these are obligately ant-associated. Patterns of association are examined from the perspective of both lycaenids and ants, with a focus on ant systematics and ecology. Lycaenids are recorded with five ant subfamilies, including the first record of an association with the Pseudomyrmecinae. The Dolichoderinae, and to some extent the Formicinae, have a disproportionately high percentage of genera that associate with lycaenid butterflies. All ant species that tend lycaenids spend at least some portion of their time foraging on vegetation to collect plant and insect nectar. There is a robust relationship between the competitive status of ants within a community, and their frequency and degree of association with lycaenids. Obligate ant-association is accompanied by a high degree of specificity for ant partner, but two notable exceptions, Ogyris aenone and O. amaryllis are discussed. Facultative myrmecophiles tend to associate with a broad range of ants, although interactions with ecologically dominant ants are less frequent than might be expected based on the abundance of dominant ant species in Australian communities.

Nectar robbers are birds, insects, or other flower visitors that remove nectar from flowers through a hole pierced or bitten in the corolla. This paper is a review of the effects of nectar robbers on pollinators, pollination, and fitness of the plants they rob. Charles Darwin assumed that nectar robbers had a negative impact on the plants that they visit, but research done in the last 50 years indicates that they often have a beneficial or neutral effect. Several studies document that robbers frequently pollinate the plants that they visit. Robbers may also have an indirect effect on the behavior of the legitimate pollinators, and in some circumstances, the change in pollinator behavior could result in improved fitness through increased pollen flow and outcrossing. The effects of nectar robbers are complex and depend, in part, on the identity of the robber, the identity of the legitimate pollinator, how much nectar the robbers remove, and the variety of floral resources available in the environment.

Plant symbiotic fungi are generally thought to express a single lifestyle that might increase (mutualism), decrease (parasitism), or have no influence (commensalism) on host fitness. However, data are presented here demonstrating that plant pathogenic Colletotrichum species are able to asymptomatically colonize plants and express nonpathogenic lifestyles.

The composition of the soil microbial community can be altered dramatically due to association with individual plant species, and these effects on the microbial community can have important feedbacks on plant ecology. Negative plant-soil feedback plays primary roles in maintaining plant community diversity, whereas positive plant-soil feedback may cause community conversion. Host-specific differentiation of the microbial community results from the trade-offs associated with overcoming plant defense and the specific benefits associated with plant rewards. Accumulation of host-specific pathogens likely generates negative feedback on the plant, while changes in the density of microbial mutualists likely generate positive feedback. However, the competitive dynamics among microbes depends on the multidimensional costs of virulence and mutualism, the fine-scale spatial structure within plant roots, and active plant allocation and localized defense. Because of this, incorporating a full view of microbial dynamics is essential to explaining the dynamics of plant-soil feedbacks and therefore plant community ecology.

Since the discovery of the effects of endophytes in the genus Acremonium on insect resistance, our understanding of these fungi has progressed greatly. However, many interactions associated with the endophytes, their hosts, and insect herbivores affect the use and expression of this resistance. This review centers on these topics, focusing on our current thinking about the use of endophyte-enhanced resistance to insects. A broader perspective is stressed in the belief that this phenomenon is only one example of many microbial-plant interactions that affect insect resistance. I begin with a review of the biology of the Acremonium endophytes, followed by a discussion of other fungal endophytes and fungal-plant interactions that result in increased insect resistance. The status of our knowledge of the effects of Acremonium endophytes on insect resistance is then presented. Finally, plant breeding, resistance management, and other influences endophytes have on agroecosystems are discussed. Background Fungal endophytes form infections of leaves and stems of healthy plants but have limited or no pathogenic effects (19). Endophytic fungi grow

The response of plants to arbuscular mycorrhizal fungi involves a temporal and spatial activation of different defence mechanisms. The activation and regulation of these defences have been proposed to play a role in the maintenance of the mutualistic status of the association, ...

Although it is well established that many insects, birds and mammals serve as important pollinators and seed dispersers of flowering plants, the role of lizards in these processes has traditionally been considered as rare and less important. However, recent work shows both that their role as mutualistic agents has been underestimated and also reveals a striking patternthat pollination and seed dispersal by lizards is most common on islands. We argue that this island phenomenon occurs because island lizards reach very high densities (density compensation) and experience a lower predation risk than do those on the mainland and, consequently, can expand their diet to include nectar, pollen and fruit. Although further empirical evidence is needed to confirm this explanation, such relationships could be ideal systems with which to study fundamental ecological problems, such as niche shifts, ecological release and competition.

The honeydew composition and production of four aphid species feeding on Tanacetum vulgare, and mutualistic relationships with the ant Lasius niger were studied. In honeydew of Metopeurum fuscoviride and Brachycaudus cardui, xylose, glucose, fructose, sucrose, maltose, melezitose, and ranose were detected. The proportion of trisaccharides (melezitose, ranose) ranged between 20% and 35%. No trisaccharides were found in honeydew of Aphis fabae, and honeydew of Macrosiphoniella tanacetaria consisted of only xylose, glucose and sucrose. M. fuscoviride produced by far the largest amounts of honeydew per time unit (880 lg/ aphid per hour), followed by B. cardui (223 lg/aphid per hour), A. fabae (133 lg/aphid per hour) and M. tanacetaria (46 lg/aphid per hour). The qualitative and quantitative honeydew production of the aphid species corresponded well with the observed attendance by L. niger. L. niger workers preferred trisaccharides over disaccharides and monosaccharides when these sugars were oered in choice tests. The results are consistent with the ants' preference for M. fuscoviride, which produced the largest amount of honeydew including a considerable proportion of the trisaccharides melezitose and ranose. The preference of L. niger for B. cardui over A. fabae, both producing similar amounts of honeydew, may be explained by the presence of trisaccharides and the higher total sugar concentration in B. cardui honeydew. M. tanacetaria, which produced only low quantities of honeydew with no trisaccharides was not attended at all by L. niger.

The animal gut serves as a primary location for the complex host-microbe interplay that is essential for homeostasis and may also reflect the types of ancient selective pressures that spawned the emergence of immunity in metazoans. In this review, we present a phylogenetic survey of gut host-microbe interactions and suggest that host defense systems arose not only to protect tissue directly from pathogenic attack but also to actively support growth of specific communities of mutualists. This functional dichotomy resulted in the evolution of immune systems much more tuned for harmonious existence with microbes than previously thought, existing as dynamic but primarily cooperative entities in the present day. We further present the protochordate Ciona intestinalis as a promising model for studying gut hostbacterial dialogue. The taxonomic position, gut physiology and experimental tractability of Ciona offer unique advantages in dissecting host-microbe interplay and can complement studies in other model systems.

To understand the evolution of insect gut symbionts it is important to determine how they are passed on to the next generation. We studied this process in Erwinia species bacteria that inhabit the gut of western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae). This is a polyphagous herbivore and a world-wide pest in agricultural crops. With bacteria in the gut, the thrips larval development time can be shorter and its oviposition rate higher compared to bacteria-free thrips. Bacteria are not directly transmitted from mother to offspring, but larvae acquire bacteria from the leaves right after they hatch. These gut bacteria are present on the leaves on feeding sites used by other thrips before the larvae arrive, probably because these other thrips have deposited bacteria via faeces or regurgitation. In this study we addressed the question whether the transmission route of symbiotic bacteria influences the thrips feeding behaviour, and determined the feeding and oviposition preference of thrips, by giving them a choice between leaves with and leaves without prior grazing by other western flower thrips. This was studied for thrips with and thrips without gut bacteria. Young larvae prefer to feed on leaves that where grazed before by other thrips and females prefer to oviposit on these grazed leaves. These results are in contradiction to earlier studies that have found that thrips larvae fitness is lower on thrips damaged plants than on clean plants. This behaviour does however promote the establishment of gut bacteria in the thrips. The factors determining the preference for thrips-damaged leaves may be the physical leaf damage or odours that are produced by the plant, the bacteria or both.

Ecological specialisation concerns all species and underlies many major ecological and evolutionary patterns.
Yet its status as a unifying concept is not always appreciated because of its similarity to concepts of the niche,
the many levels of biological phenomena to which it applies, and the complexity of the mechanisms influencing
it. The evolution of specialisation requires the coupling of constraints on adaptive evolution with covariation
of genotype and environmental performance. This covariation itself depends upon organismal properties
such as dispersal behaviour and life history and complexity in the environment stemming from factors such
as species interactions and spatio-temporal heterogeneity in resources. Here, we develop a view on
specialisation that integrates across the range of biological phenomena with the goal of developing a more
predictive conceptual framework that specifically accounts for the importance of biotic complexity and
coevolutionary events.

All plants studied in natural ecosystems are symbiotic with fungi that either reside entirely (endophytes) or partially (mycorrhizae) within plants. These symbioses appear to adapt to biotic and abiotic stresses and may be responsible for the survival of both plant hosts and fungal symbionts in high stress habitats. Here we describe the role of symbiotic fungi in plant stress tolerance and present a strategy based on adaptive symbiosis to potentially mitigate the impacts of global change on plant communities.

A central question in behavioral ecology has been why animals live in groups. Previous theories about the evolution of sociality focused on the potential benefits of decreased risk of predation, increased foraging or feeding efficiency, and mutual aid in defending resources and/or rearing offspring. This paper argues that access to mutualistic endosymbiotic microbes is an underappreciated benefit of group living and sets out to reinvigorate Troyer's hypothesis that the need to obtain cellulolytic microbes from conspecifics influenced the evolution of social behavior in herbivores and to extend it to nonherbivores. This extension is necessary because the benefits of endosymbionts are not limited to nutrition; endosymbionts also help protect their hosts from pathogens. When hosts must obtain endosymbionts from conspecifics, they are forced to interact. Thus, complex forms of sociality may be more likely to evolve when hosts must repeatedly obtain endosymbionts from conspecifics than when endosymbionts can be obtained either directly from the environment, are vertically transmitted, or when repeated inoculations are not necessary. Observations from a variety of taxa are consistent with the ideas that individuals benefit from group living by gaining access to endosymbionts and the complexity of social behavior is associated with the mode of acquisition of endosymbionts. Ways to test this theory include (a) experiments designed to examine the effects of endosymbionts on host fitness and how endosymbionts are obtained and (b) using phylogenetic analyses to examine endosymbiont-host coevolution with the goal of determining the relationship between the mode of endosymbiont acquisition and host sociality.

Mycorrhizal phenotypes arise from interactions among plant and fungal genotypes and the environment. Differences in the stoichiometry and uptake capacity of fungi and plants make arbuscular mycorrhizal (AM) fungi inherently more nitrogen (N) limited and less phosphorus (P) limited than their host plants. Mutualistic phenotypes are most likely in P-limited systems and commensal or parasitic phenotypes in N-limited systems. Carbon (C) limitation is expected to cause phenotypes to shift from mutualism to commensalism and even parasitism. Two experiments compared the influence of fertilizer and shade on mycorrhizas in Andropogon gerardii across three naturally N-limited or P-limited grasslands. A third experiment examined the interactive effects of N and P enrichment and shade on A. gerardii mycorrhizas. Our experiments generated the full spectrum of mycorrhizal phenotypes. These findings support the hypothesis that mutualism is likely in P-limited systems and commensalism or parasitism...

Phytochrome-mediated perception of the ratio of red to far-red wavelengths in the ambient light environment is fundamental to plant growth and development. Such monitoring enables plants to detect neighboring vegetation and initiate avoidance responses, thus conferring considerable selective advantage. The shade avoidance syndrome in plants is characterized by elongation growth and early flowering, responses that are fully induced by end-of-day far-red light treatments. Elucidating the roles of individual phytochromes in mediating responses to red to far-red has however always been confounded by synergistic and mutually antagonistic coactions between family members. The creation of triple and quadruple mutants in Arabidopsis, deficient in multiple phytochromes, has revealed functional redundancy between phyB, D, and E in controlling flowering time, leaf development, and regulation of the homeobox gene, ATHB-2. In addition, mutant analysis suggests a possible novel role for phyC in suppressing ATHB-2 transcription in the light. fax 44 -0116 -252-3330.

Key advances are being made on the structures of predator-prey food webs and competitive communities that enhance their stability, but little attention has been given to such complexity-stability relationships for mutualistic communities. We show, by way of theoretical analyses with empirically informed parameters, that structural properties can alter the stability of mutualistic communities characterized by nonlinear functional responses among the interacting species. Specifically, community resilience is enhanced by increasing community size (species diversity) and the number of species interactions (connectivity), and through strong, symmetric interaction strengths of highly nested networks. As a result, mutualistic communities show largely positive complexity-stability relationships, in opposition to the standard paradox. Thus, contrary to the commonlyheld belief that mutualismÕs positive feedback destabilizes food webs, our results suggest that interplay between the structure and function of ecological networks in general, and consideration of mutualistic interactions in particular, may be key to understanding complexity-stability relationships of biological communities as a whole.

Not all floral visitors attracted to flowers are pollinators. Instead, some visitors circumvent the floral opening, usually removing nectar without contacting the anthers and/or stigma. Here we review the evolutionary ecology of nectar robbing from both the plant and animal perspective. Effects of robbing on female and male components of plant reproduction range from negative to positive. Their underlying mechanisms are diverse, including direct effects and indirect effects mediated through changes in pollination. We detail how plants may be able to deter robbers through morphological and chemical traits. For the evolutionary ecology of robbing to move beyond a phytocentric perspective, studies must also address the causes of robbing and the consequences for both robbers and pollinators. We use an energetics approach to evaluate these causes and consequences. Finally, we highlight unanswered questions in need of further research.

The idea that phytopathogenic fungi associated with tree-killing bark beetles are critical for overwhelming tree defenses and incurring host tree mortality, herein called the classic paradigm (CP), has driven research on bark beetle-fungus symbiosis for decades. It has also strongly influenced our views of bark beetle ecology. We discuss fundamental flaws in the CP, including the lack of consistency of virulent fungal associates with tree-killing bark beetles, the lack of correspondence between fungal growth in the host tree and the development of symptoms associated with a successful attack, and the ubiquity of similar associations of fungi with bark beetles that do not kill trees. We suggest that, rather than playing a supporting role for the host beetle (tree killing), phytopathogenicity performs an important role for the fungi. In particular, phytopathogenicity may mediate competitive interactions among fungi and support survival and efficient resource capture in living, defensive trees.

We discuss studies of foliar endophytic fungi (FEF) and arbuscular mycorrhizal fungi (AMF) associated with Theobroma cacao in Panama. Direct, experimentally controlled comparisons of endophyte free (EÀ) and endophyte containing (Eþ) plant tissues in T. cacao show that foliar endophytes (FEF) that commonly occur in healthy host leaves enhance host defenses against foliar damage due to the pathogen (Phytophthora palmivora). Similarly, root inoculations with commonly occurring AMF also reduce foliar damage due to the same pathogen. These results suggest that endophytic fungi can play a potentially important mutualistic role by augmenting host defensive responses against pathogens. There are two broad classes of potential mechanisms by which endophytes could contribute to host protection: (1) inducing or increasing the expression of intrinsic host defense mechanisms and (2) providing additional sources of defense, extrinsic to those of the host (e.g., endophytebased chemical antibiosis). The degree to which either of these mechanisms predominates holds distinct consequences for the evolutionary ecology of host-endophyte-pathogen relationships. More generally, the growing recognition that plants are composed of a mosaic of plant and fungal tissues holds a series of implications for the study of plant defense, physiology, and genetics.

The rhizosphere sediments of seagrasses are generally a site of intense nitrogen fixation activity and this can provide a significant source of``new'' nitrogen for the growth of the plants. In this paper, I review the data concerning nitrogen fixation in seagrass ecosystems, the transfer of the fixed nitrogen from the bacteria to the plants and its contribution to the overall productivity of seagrasses in different climatic zones. The relationship between the plants and diazotrophic heterotrophic bacteria in the rhizosphere is discussed, particularly focusing on the potentially important role of nitrogen-fixing, sulphate-reducing bacteria. The regulation of nitrogen fixation rates in the rhizosphere by photosynthetically driven oxygen and fixed carbon release by the plant roots and rhizomes, and the availability of ammonium in the porewater, is assessed. Finally, the hypothesis that a mutualistic or symbiotic association exists between the seagrasses and heterotrophic nitrogen fixers in the rhizosphere, based on the mutual exchange of fixed carbon and nitrogen, is discussed.

We tested whether a plant's life time seed production is increased by parasitization of herbivores in a tritrophic system, Arabidopsis thaliana (Brassicaceae) plants, Pieris rapae (Lepidoptera: Pieridae) caterpillars and the solitary endoparasitoid Cotesia rubecula (Hymenoptera: Braconidae). We established seed production for intact A. thaliana plants, plants that were mechanically damaged, plants fed upon by parasitized caterpillars and plants fed upon by unparasitized caterpillars. In the first experiment, with ecotype Landsberg (erecta mutant), herbivory by unparasitized P. rapae caterpillars resulted in a strongly reduced seed production compared to undamaged plants. In contrast, damage by P. rapae caterpillars that had been parasitized by C. rubecula did not result in a significant reduction in seed production. For the second experiment with the ecotype Columbia, the results were identical. Plants damaged by unparasitized caterpillars only produced seeds on regrown shoots. Seed production of plants that had been mechanically damaged was statistically similar to that of undamaged plants. Production of the first ripe siliques by plants fed upon by unparasitized caterpillars was delayed by 18-22 days for Landsberg and 9-10 days for Columbia. We conclude that parasitization of P. rapae by C. rubecula potentially confers a considerable fitness benefit for A. thaliana plants when compared to plants exposed to feeding damage by unparasitized P. rapae larvae. Plants that attract parasitoids and parasitoids that respond to herbivore-induced plant volatiles will both experience selective advantage, justifying the use of the term mutualism for this parasitoid-plant interaction. This type of mutualism is undoubtedly very common in nature.

Background: Attine ants live in an intensely studied tripartite mutualism with the fungus Leucoagaricus gongylophorus, which provides food to the ants, and with antibiotic-producing actinomycete bacteria. One hypothesis suggests that bacteria from the genus Pseudonocardia are the sole, co-evolved mutualists of attine ants and are transmitted vertically by the queens. A recent study identified a Pseudonocardia-produced antifungal, named dentigerumycin, associated with the lower attine Apterostigma dentigerum consistent with the idea that co- evolved Pseudonocardia make novel antibiotics. An alternative possibility is that attine ants sample actinomycete bacteria from the soil, selecting and maintaining those species that make useful antibiotics. Consistent with this idea, a Streptomyces species associated with the higher attine Acromyrmex octospinosus was recently shown to produce the well-known antifungal candicidin. Candicidin production is widespread in environmental isolates of Streptomyces, so this could either be an environmental contaminant or evidence of recruitment of useful actinomycetes from the environment. It should be noted that the two possibilities for actinomycete acquisition are not necessarily mutually exclusive.
Results: In order to test these possibilities we isolated bacteria from a geographically distinct population of A. octospinosus and identified a candicidin-producing Streptomyces species, which suggests that they are common mutualists of attine ants, most probably recruited from the environment. We also identified a Pseudonocardia species in the same ant colony that produces an unusual polyene antifungal, providing evidence for co-evolution of Pseudonocardia with A. octospinosus.
Conclusions: Our results show that a combination of co-evolution and environmental sampling results in the diversity of actinomycete symbionts and antibiotics associated with attine ants.

Non-Darwinian theories about the emergence and evolution of complexity date back at least to Lamarck, and include those of Herbert Spencer and the "emergent evolution" theorists of the later nineteenth and early twentieth centuries. In recent decades, this approach has mostly been espoused by various practitioners in biophysics and complexity theory. However, there is a Darwinian alternative - in essence, an economic theory of complexity - proposing that synergistic effects of various kinds have played an important causal role in the evolution of complexity, especially in the "major transitions". This theory is called the "synergism hypothesis". We posit that otherwise unattainable functional advantages arising from various cooperative phenomena have been favored over time in a dynamic that the late John Maynard Smith characterized and modeled as "synergistic selection". The term highlights the fact that synergistic "wholes" may beco...