In situ tests on inducible defenses in Dictyota kunthii and Macrocystis integrifolia (Phaeophyceae) from the Chilean coast

Numerous experimental studies have reported inducible defenses in macroalgae, but most of them have been conducted in laboratory environments where algae were maintained detached from the substratum and in artificial flow regimes. The results of those experiments might not reflect the natural situation, which can only be studied in situ. We examined whether the brown macroalgae Dictyota kunthii (C. Agardh) Greville and Macrocystis integrifolia (Bory) show inducible defenses following exposure to different grazing levels (direct, water-borne cues from nearby grazed conspecifics, presence of a non-grazing herbivore and natural grazing) in field experiments, striving to maintain natural conditions as much as possible. We measured palatability of algae after exposure to different grazing levels by using live algae and agar-based food containing non-polar extracts. M. integrifolia showed no induction of defenses (at least not of non-polar compounds), suggesting constitutive defenses, absence of defenses (tolerance) or use of another strategy to avoid herbivory. These results are similar to those from previous laboratory experiments. In D. kunthii, defense was induced after two weeks of direct grazing by amphipods under field conditions. Water-borne cues from nearby grazed conspecifics, presence of a non-grazing herbivore and natural grazing did not induce defenses. Induction of defense in response to direct grazing agrees with results from a previous laboratory study, but while indirect cues induced defenses in the laboratory, there was no measurable induced defense in the field. Probably chemical cues from grazers are diluted quickly in the field, not reaching concentrations that cause induction of defenses. This might be the reason why in some algae induction by direct grazing is a more important defensive strategy than induction by water-borne cues. The results from our study also suggest that laboratory experiments showing induced defenses in response to grazed neighbours or mere grazer presence need to be interpreted with caution.     

Testing for the induction of anti-herbivory defences in four Portuguese macroalgae by direct and water-borne cues of grazing amphipods

Herbivory is a key factor in regulating plant biomass, thereby driving ecosystem performance. Algae have developed multiple adaptations to cope with grazers, including morphological and chemical defences. In a series of experiments we investigated whether several species of macroalgae possess anti-herbivore defences and whether these could be regulated to demand, i.e. grazing events. The potential of direct grazing on defence induction was assessed for two brown (Dictyopteris membranacea, Fucus vesiculosus) and two red seaweeds (Gelidium sesquipedale, Sphaerococcus coronopifolius) from São Rafael and Ria Formosa, Portugal. Bioassays conducted with live algal pieces and agar-based food containing lipophilic algal extracts were used to detect changes in palatability after exposure to amphipod attacks (=treatment phase). Fucus vesiculosus was the only species significantly reducing palatability in response to direct amphipod-attacks. This pattern was observed in live F. vesiculosus pieces and agar-based food containing a lipophilic extract, suggesting that lipophilic compounds produced during the treatment phase were responsible for the repulsion of grazers. Water-borne cues of grazed F. vesiculosus as well as non-grazing amphipods also reduced palatability of neighbouring conspecifics. However, this effect was only observed in live tissues of F. vesiculosus. This study is the first to show that amphipods, like isopods, are capable to induce anti-herbivory defences in F. vesiculosus and that a seasonally variable effectiveness of chemical defences might serve as a dynamic control in alga–herbivore interactions.     

Effect of Detachment on the Palatability of two Kelp Species

Many species of macroalgae survive after becoming dislodged from their primary substratum, but little is known about their capacity to express anti-herbivore defences after detachment. We examined the effect of detachment on the relative palatability of the two kelp species Lessonia nigrescens and Macrocystis integrifolia to mesograzers. Laboratory and field experiments were conducted on the northern-central coast of Chile to investigate whether (i) time after detachment and (ii) grazing on detached and attached algae could trigger internal defence mechanisms in the algae, which may have acted as deterrents to grazing. In order to examine palatability, feeding assays were run after each experiment using fresh algal pieces and artificial food. Time after detachment had a significant influence on palatability of L. nigrescens but not of M. integrifolia. During the first 12 days of detachment, detached L. nigrescens held in grazer-free laboratory tanks were not significantly more palatable than attached conspecifics from the field but thereafter detached individuals became more palatable. Floating individuals of M. integrifolia showed no effect of detachment, indicating that this alga maintains its defence after detachment. An experiment conducted in the field confirmed these results for M. integrifolia. An additional laboratory experiment confirmed that attachment status plays an important role on algal defence reaction for L. nigrescens when exposed to grazers. Detached and previously grazed individuals of this species were less palatable than grazer-free control algae, but grazing had no effect on palatability of attached algae. Our results indicate that kelps have varying capacities for development of anti-grazing responses once they become detached, possibly depending on their capacity to float and survive after detachment.     

Inducible defence and its modulation by environmental stress in the red alga Chondrus yendoi (Yamada and Mikami in Mikami, 1965) from Honshu Island, Japan

Among the numerous anti-herbivore defences developed by macroalgae, chemical and morphological traits are best documented and understood. Plant defence theory suggests that these resistances, which can either be constitutive or inducible, are associated with metabolic costs. They should therefore be impaired under conditions of energy limitation, but evidence for this prediction is scarce. In two subsequent experiments, we tested whether a reduction of light availability is changing feeding rates of the two mesoherbivores Idotea ochotensis and Lacuna smithii on the red alga Chondrus yendoi. Algal individuals were kept in outdoor mesocosm facilities for 10 days, during which we manipulated the amount of incoming sunlight at 6 levels (0%-99% reduction, i.e. 2000-20 μmol s^− 1 m^− 2). Orthogonal to this, we established the presence or absence of one of the herbivores to test whether C. yendoi can generate a defence. Algal palatability was investigated afterwards in no-choice feeding assays using naïve grazer individuals. The consumption of algal tissue in L. smithii increased with decreasing light, while this was not the case for I. ochotensis. However, we found a defence induced as a reaction to herbivory only by the highly mobile isopod but not when the slowly moving snail grazed on C. yendoi. Isopod total consumption rates in our experiments were 40 times higher than those of the gastropods. We therefore suggest that C. yendoi exhibits grazer-specific reactions to herbivory, depending on the mobility and voracity of the consumers. Interestingly, only for one of the grazers, i.e. the snail, short-term light reduction influenced the palatability of algal tissue. We discuss different but not mutually exclusive models that could explain this pattern. In conclusion, we view this three-species system as an illustrative example for specificity in grazer-algal interactions and their modification by environmental stress.     

INDUCTION AND REDUCTION OF ANTI‐HERBIVORE DEFENSES IN BROWN AND RED MACROALGAE OFF THE KENYAN COAST1

Herbivory is particularly intense in tropical benthic communities, suggesting preference of constitutive, rather than inducible, anti‐herbivory defense. The objective of the study was to examine whether anti‐herbivore defenses in the red alga Hypnea pannosa J. Agardh and the brown algae Sargassum asperifolium Hering and G. Martens ex J. Agardh and Cystoseira myrica (S.G. Gmelin) C. Agardh could be induced and subsequently reduced in response to grazing by the amphipod Cymadusa filosa Savigny. During a 14‐day treatment phase, algae were exposed to amphipod grazing or were left ungrazed (control). Subsequently, one subset of algae was used in feeding assays, whereas another was cultivated for additional 14 days without consumers (recovery phase). At the end of each phase, bioassays were conducted to detect defensive traits in terms of differences in consumption rates of grazed and control pieces of live algae and agar‐based food containing nonpolar algal extracts. Consumption of grazed live S. asperifolium and H. pannosa specimens was lower than of control algae. Furthermore, nonpolar extracts of grazed S. asperifolium and C. myrica were less preferred than those from control algae. Defensive responses were exclusively detected after the treatment phase, although strong preference of ungrazed H. pannosa and C. myrica over grazed conspecifics continued throughout the recovery phase. These findings suggest that phenotypic plasticity in anti‐herbivory defense of marine macroalgae 1) might be more common than previously shown, 2) could be switched on and off within 2 weeks, and 3) can be found in nonpolar algal extracts.     

Effects of Chiton granosus (Frembly, 1827) and other molluscan grazers on algal succession in wave exposed mid-intertidal rocky shores of central Chile

Molluscan grazers can have important effects on the abundance, colonization rates, and successional pathways of algal assemblages and the entire intertidal community. In general, early successional algae are more readily consumed than corticated algae and kelps, which usually get established later in the community succession. To generalize, however, the effect of different grazers on algal assemblages must be examined on different coasts and under different scenarios. This information could help us understand the mechanisms of ecosystem processes and situations in which general models do not apply. Along the coast of Chile, humans harvest large keyhole limpets, which seem to be the only invertebrate grazers capable of controlling the dominant corticated alga Mazzaella laminarioides, a canopy-forming species that can cover extensive areas of the mid intertidal zone. In this scenario, where large limpets are harvested, the overall effects of the diverse molluscan assemblage of limpets, chitons and snails on algal succession and on corticated algae in particular are not clear. We conducted a 26-month-long experiment to evaluate the effects of molluscan grazers on mid-intertidal algal succession and to isolate the effects of Chiton granosus, the most conspicuous member of the assemblage at these tidal elevations. At sites heavily impacted by humans the molluscan grazer assemblage had strong negative effects on colonization and abundance of green algae such as ulvoids and Blidingia minima. In doing so, the grazer assemblage had a strong negative indirect effect on the establishments of chironomid fly larvae, which were only observed on green algal mats and rarely on bare rock. No significant effects were detected on epilithic microalgae, and effects on sessile invertebrates were highly variable over space and time. C. granosus also had significant negative effects on green algae but did not account for the total grazing pressure exerted by the guild. Limited foraging excursions (ca. 35 cm) from refuges and moderate site (crevice) fidelity in this species may contribute to the patchiness in green algal distribution observed in the field. Nearly 13 months after rock surface were experimentally cleared, M. laminarioides appeared in all experimental plots, but increased over three times faster in enclosures containing C. granosus than in exclosures plots or controls, suggesting that moderate levels of herbivory could actually facilitate the establishment of this alga in the succession and that the green algal cover found in the absence of grazers may delay its establishment.     

Ecological and Evolutionary Interactions between Wild Crucifers and Their Herbivorous Insects

Abstract Insects feeding on ten species of wild crucifer were investigated. Differences in host plant range and insect community structure were examined with regard to anti-herbivore defense mechanisms. Most of the crucifer species deterred insect herbivory by disappearing in the summer or by lowering their intrinsic quality as food for insects. Species with these defense mechanisms were exploited by only a few specialized herbivorous insects that seemed to have counter defenses. The plants without these defense mechanisms were used by many herbivorous insect species. Rorippa indica lacked direct defenses, but supported a low total density of herbivore individuals. This crucifer has an indirect defense mechanism: ants attracted to floral nectar defended the plant from deleterious herbivores. Crucifers that disappeared seasonally lacked other anti-herbivore defense mechanisms. This suggests that the phonological response is an alternative other responses to herbivore attack.     

Intra- and interspecific differences in palatability of Arctic macroalgae from Kongsfjorden (Spitsbergen) for two benthic sympatric invertebrates

Information on herbivore-macroalgae interactions is extremely scarce for Arctic habitats. In this study, the potential of 19 macroalgae as food source for herbivores was investigated for the first time in Arctic waters (Kongsfjorden, Spitsbergen) with emphasis on algal defense against grazing. Only two of the 19 tested macroalgae-associated invertebrates consumed macroalgae in measurable amounts, the locally abundant green sea urchin Strongylocentrotus droebachiensis (OF Müller) and the amphipod Gammarellus homari (JC Fabricius). In the inner basin of the fjord, the dense macroalgal stock represents a potential food source. However, in this area, herbivory plays only a minor role. In contrast, in the outer basin of Kongsfjorden S. droebachiensis exerts a strong top-down control on macroalgal assemblages.Laboratory feeding assays in the with the two herbivores showed grazer-specific feeding preferences. The amphipod G. homari exhibits a preference for delicate red algal species like Devaleraea ramentacea, whereas the sea urchin S. droebachiensis significantly preferred more leathery seaweeds like Laminaria and Alaria. The red alga Palmaria palmata is a very attractive food for both herbivores, while the brown alga Desmarestia viridis and the red alga Ptilota gunneri are among the least preferred algae. To distinguish between physical and tissue-specific plant properties, which have a deterring or stimulating effect on the grazing behaviour, both grazers were offered intact algal tissue and artificial food in separate feeding assays. While physical and tissue-specific plant properties-especially in the Laminariales-deterred G. homari, these properties did not deter S. droebachiensis.This study provides insights into herbivore consumption in the benthic food web of Kongsfjorden. As a general potential top-down factor controlling primary production, herbivory needs to be quantified further in Kongsfjorden to develop adequate carbon flow models for this important reference site for the study of climate change on high latitude marine ecosystems.     

Resistance of the brown alga Fucus vesiculosus to herbivory

Herbivory is typically intense in marine littoral environments; thus, macrophytes are expected to evolve defenses against grazing. Although putative defenses of macrophytes are widely studied, there is lack of studies demonstrating the main premises of defense adaptations: the consequences of herbivory to macrophytes, genetic variation of defense traits and the costs and benefits of defenses in natural environment. We conducted a factorial experiment, where we manipulated amount of herbivory, growing depth and nutrient availability, and measured resistance to herbivory as well as genetic variation and costs of phlorotannins, putative defensive secondary metabolites, in the brown alga Fucus vesiculosus . Herbivory on algae varied with depth: grazing did not cause losses close to the surface, but, most of the algal production was consumed at the deeper end of the algal belt. The higher the genotypic phlorotannin content the less damage the genotype received implying that phlorotannins acted as a resistance trait. Production of phlorotannins was associated with costs for growth. Consistent with the prediction that the cost of defense will be greatest when resources are limiting, the cost appeared only in the deep end of the algal belt where growth was slowed down. Phlorotannins displayed phenotypic plasticity; the three factors influenced phlorotannins interactively, with the main tendencies of nutrient enrichment decreasing and herbivory and increasing depth increasing phlorotannins. Despite this plasticity, variation of phlorotannins was mainly due to the genotype of algae. These results emphasize the role of herbivory as a selective agent for algal defenses and the importance of genetic variation in the constitutive level of phlorotannins in interactions with natural enemies. The cost of phlorotannins may constrain the evolution of resistance in environments where growth is limited by light availability.     

Grazers and Phytoplankton Growth in the Oceans: an Experimental and Evolutionary Perspective

The taxonomic composition of phytoplankton responsible for primary production on continental shelves has changed episodically through Earth history. Geological correlations suggest that major changes in phytoplankton composition correspond in time to changes in grazing and seawater chemistry. Testing hypotheses that arise from these correlations requires experimentation, and so we carried out a series of experiments in which selected phytoplankton species were grown in treatments that differed with respect to the presence or absence of grazers as well as seawater chemistry. Both protistan (Euplotes sp.) and microarthropod (Acartia tonsa) grazers changed the growth dynamics and biochemical composition of the green alga Tetraselmis suecica, the diatom Thalassiosira weissflogii, and the cyanobacterium Synechococcus sp., increasing the specific growth rate and palatability of the eukaryotic algae, while decreasing or leaving unchanged both parameters in the cyanobacteria. Synechococcus (especially) and Thalassiosira produced toxins effective against the copepod, but ciliate growth was unaffected. Acartia induced a 4-6 fold increase of Si cell quota in the diatom, but Euplotes had no similar effect. The differential growth responses of the eukaryotic algae and cyanobacteria to ciliate grazing may help to explain the apparently coeval radiation of eukaryophagic protists and rise of eukaryotes to ecological prominence as primary producers in Neoproterozoic oceans. The experimental results suggest that phytoplankton responses to the later radiation of microarthropod grazers were clade-specific, and included changes in growth dynamics, toxin synthesis, encystment, and (in diatoms) enhanced Si uptake.     

Algal propagule banks modify competition, consumer and resource control on Baltic rocky shores

We hypothesized that supply from macroalgal propagule banks may influence the relative abundance of annual and perennial algae and that this may alter the effects of grazers and nutrients on species composition. In a factorial field experiment in the Baltic Sea littoral system we tested the effects of manipulating propagule banks, the abundance of crustacean and gastropod grazers, and nutrient supply on recruitment and growth of macroalgae over a year. Moreover, we determined seasonal patterns of macroalgal propagule dispersal at the experimental site and quantified algal abundance and recruitment at 25 locations throughout the Baltic Sea. Experimental manipulations had minor effects on adults of the dominating perennial alga, Fucus vesiculosus. Instead, we found that species composition was determined by processes operating at early life stages. Propagule supply from a propagule bank strongly favored the fast-growing annual alga Enteromorpha spp. which then blocked settlement and recruitment of Fucus. Grazers reduced the abundance of annual algae and indirectly favored Fucus recruitment. There was an apparent trade-off between gains from the propagule bank and losses to herbivory in five of seven colonizing species. Nutrient enrichment overrode grazer control of annual algae and accelerated the decline of Fucus only when annual algae had already achieved high densities through the propagule bank. Corroborating the experimental findings, field surveys across the Baltic showed that Fucus recruit densities can be predicted from the cover of annual algae during the period of Fucus reproduction and settlement. Recruitment inhibition by annual algae, which is driven by the abundance of annuals in the propagule bank, increasing nutrient levels, and declining consumer control, is suggested as a mechanistic explanation of the current decline of perennial algae in the Baltic Sea.     

Preliminary survey of toxicity of the green algaCaulerpa taxifolia introduced into the Mediterranean

Caulerpa taxifolia (Vahl) C. Agardh (Ulvophyceae, Caulerpales) is an alga of tropical origin that was accidentally introduced into the Mediterranean sea in 1984, where this species can reach an abundance that has never been described in tropical endemic regions. It is known that caulerpacean algae can develop an efficient strategy against grazers consisting of the synthesis of repulsive of toxic secondary metabolites: we report here the first study of the toxicity of purified secondary metabolites and raw extracts fromC. taxifolia from the Mediterranean.Toxicity was evaluated on three models: mice (lethality), mammalian cells in culture (cytotoxicity) and sea urchin eggs (disturbance of cell proliferation). Aqueous extracts are only active on fibroblasts and mice. In the three toxicity models a seasonal variation of toxicity is observed for the crude methanol extract as well as a decrease of this activity whenC. taxifolia from the Mediterranean is kept in aquaria. Pure compounds exhibit different toxicity depending on the assay. 10,11-epoxycaulerpenyne is the most active substance on mice and fibroblasts whereas taxifolial A and D are inactive or only weakly toxic. Among the four tested compounds caulerpenyne, the major metabolite ofC. taxifolia, is the most active on sea urchin eggs. Caulerpenyne may therefore represent an ecological risk for microorganisms and the eggs of multicellular animals living close to this alga. The ecological impact of this toxicity on marine organisms and the interaction of this alga with the herbivorous fauna are discussed.     

Compensatory growth of the kelp Macrocystis integrifolia (Phaeophyceae, Laminariales) against grazing of Peramphithoe femorata (Amphipoda, Ampithoidae) in northern-central Chile

Compensation of tissue loss has been considered an alternative strategy for seaweeds that have no or only minor chemical or structural defense against herbivory. Compensatory responses are facilitated by resource transfer among different tissues and have been suggested for large kelps. Macrocystis integrifolia (Bory) is a common kelp species from northern-central Chile, which is characterized by high growth rates and the absence of lipophilic chemical defenses against herbivore grazing. Herein, we used the giant kelp M. integrifolia to test for compensatory growth in response to grazing by the nest-dwelling amphipod Peramphithoe femorata (Krøyer). Amphipods were allowed to graze inside nests on subapical blades of M. integrifolia sporophytes for 14 days. We measured growth and chemical composition (C, N, laminaran and mannitol) of apical and subapical blades of grazed and ungrazed (control) sporophytes. Our results revealed the capability of M. integrifolia to maintain elongation rates in grazed subapical blades, which were similar to those of subapical blades from ungrazed sporophytes. Apical blades grew slower in grazed than in ungrazed sporophytes indicating a trade-off between apical and subapical blades when herbivores were present. Thus, compensation occurs in blades directly attacked by grazers and is probably mediated by vertical resource allocation within sporophytes to subapical blades, a suggestion supported by the fact that stipe internodes in these regions grew more on grazed sporophytes. In general, our study indicates that M. integrifolia exhibits compensatory growth against the herbivore amphipod P. femorata, and we suggest that this could be an important strategy of large kelp species to tolerate moderate grazing intensities.     

Expression and costs of induced defense traits in Alliaria petiolata,a widespread invasive plant

We used jasmonic acid to induce first-year plants of Alliaria petiolata, a European invader that largely escapes herbivory in North America, to examine continental, population, and environmental variation in the expression and costs of induced defense traits. While absolute levels varied among populations, the induction of trypsin inhibitor activity was strong and largely uniform across five native and seven invasive populations. Trichome densities varied across populations, were absent in two of them, and only tended to be inducible by jasmonic acid. Jasmonate induction was substantially costly to leaf growth and dry biomass production, the magnitude of which varied little among populations. Continental origin of the populations explained an insignificant amount of variation in any trait. Trypsin inhibitor activity was strongly inducible across a nutrient gradient, but induction was more costly to leaf growth at low soil nutrient levels. Our results show that A. petiolata displays defense traits that are strongly inducible by jasmonic acid across populations, that jasmonate induction is substantially costly to growth with little variation among populations, and that costs of induction increase with decreased soil nutrient availability. Escaping the need to express induced defense traits and their costs in the face of reduced herbivory in introduced habitats may benefit fitness of invasive plants even in the absence of any evolutionary change in resistance in these plants.     

Direct and indirect effects of ocean acidification and warming on a marine plant–herbivore interaction

The impacts of climatic change on organisms depend on the interaction of multiple stressors and how these may affect the interactions among species. Consumer–prey relationships may be altered by changes to the abundance of either species, or by changes to the per capita interaction strength among species. To examine the effects of multiple stressors on a species interaction, we test the direct, interactive effects of ocean warming and lowered pH on an abundant marine herbivore (the amphipod Peramphithoe parmerong), and whether this herbivore is affected indirectly by these stressors altering the palatability of its algal food (Sargassum linearifolium). Both increased temperature and lowered pH independently reduced amphipod survival and growth, with the impacts of temperature outweighing those associated with reduced pH. Amphipods were further affected indirectly by changes to the palatability of their food source. The temperature and pH conditions in which algae were grown interacted to affect algal palatability, with acidified conditions only affecting feeding rates when algae were also grown at elevated temperatures. Feeding rates were largely unaffected by the conditions faced by the herbivore while feeding. These results indicate that, in addition to the direct effects on herbivore abundance, climatic stressors will affect the strength of plant–herbivore interactions by changes to the susceptibility of plant tissues to herbivory.     

Patterns in shallow subtidal marine assemblages along the coast of New South Wales

Shallow subtidal areas of rocky reefs in central and southern New South Wales may best be described as a mosaic of habitats, the distributions of which are seemingly related to depth, wave exposure and a number of biological processes, particularly herbivory. The Fringe habitat is generally found only in the most shallow waters. Forests of the laminarian alga Ecklonia radiata are often found at intermediate depths. In deeper, or more sheltered water, sponges, ascidians and red algae are more abundant and the abundances of sea urchins and other invertebrate grazers decline. Overlying this broad-brush pattern are patches of crustose coralline algae (the Barrens habitat), the distributions of which are not clearly related to depth. Invertebrate herbivores, and sea urchins in particular, are abundant in the Barrens habitat. The Barrens habitat was most represented at the more southern locations. At the two most northern locations, reefs were shorter in length and dominated by ascidians (Pyura species).     

Metabolomic assessment of induced and activated chemical defence in the invasive red alga Gracilaria vermiculophylla

In comparison with terrestrial plants the mechanistic knowledge of chemical defences is poor for marine macroalgae. This restricts our understanding in the chemically mediated interactions that take place between algae and other organisms. Technical advances such as metabolomics, however, enable new approaches towards the characterisation of the chemically mediated interactions of organisms with their environment. We address defence responses in the red alga Gracilaria vermiculophylla using mass spectrometry based metabolomics in combination with bioassays. Being invasive in the north Atlantic this alga is likely to possess chemical defences according to the prediction that well-defended exotics are most likely to become successful invaders in systems dominated by generalist grazers, such as marine macroalgal communities. We investigated the effect of intense herbivore feeding and simulated herbivory by mechanical wounding of the algae. Both processes led to similar changes in the metabolic profile. Feeding experiments with the generalist isopod grazer Idotea baltica showed that mechanical wounding caused a significant increase in grazer resistance. Structure elucidation of the metabolites of which some were up-regulated more than 100 times in the wounded tissue, revealed known and novel eicosanoids as major components. Among these were prostaglandins, hydroxylated fatty acids and arachidonic acid derived conjugated lactones. Bioassays with pure metabolites showed that these eicosanoids are part of the innate defence system of macroalgae, similarly to animal systems. In accordance with an induced defence mechanism application of extracts from wounded tissue caused a significant increase in grazer resistance and the up-regulation of other pathways than in the activated defence. Thus, this study suggests that G. vermiculophylla chemically deters herbivory by two lines of defence, a rapid wound-activated process followed by a slower inducible defence. By unravelling involved pathways using metabolomics this work contributes significantly to the understanding of activated and inducible defences for marine macroalgae.     

Effects of competition and herbivory on interactions between a hard coral and a brown alga

Despite widespread acceptance of the negative effects of macroalgae on corals, very few studies have experimentally tested the competitive nature of the interaction, and most have ignored the potential effects of corals on algae. We report the effects of herbivory and competition on the growth of the branching scleractinian coral Porites cylindrica Dana and the creeping foliose brown alga Lobophora variegata (Lamouroux) Womersley, on an inshore fringing reef of the central Great Barrier Reef. L. variegata overgrows branches of P. cylindrica from the base up, forming a distinct boundary between the alga and the coral tissue. The experiment used exclusion cages to test for effects of herbivores, and removal of algae and coral tissue, at their interaction boundary, to test for inhibition of the competitors by each other. Comparisons of coral branches with the algae present or removed showed that the presence and overgrowth of the alga caused significant coral tissue mortality. Comparisons of branches with coral tissue unmanipulated or damaged showed that the coral inhibited the overgrowth by L. variegata, but that the algae were markedly superior competitors. Importantly, reduced herbivory resulted in faster algal growth and consequent overgrowth and mortality of coral tissue, demonstrating the critical importance of herbivory to the outcome of the competitive interaction.     

The Potential Anti-Herbivory Role of Microorganisms on Plant Thorns

Thorns, spines and prickles are some of the anti-herbivore defenses that plants have evolved. They were recently found to be commonly aposematic (warning coloration). However, the physical anti-herbivore defense executed by these sharp structures seems to be only the tip of the iceberg. We show that thorns of various plant species commonly harbor an array of aerobic and anaerobic pathogenic bacteria including Clostridium perfringens the causative agent of the life-threatening gas gangrene, Bacillus anthracis, and Pantoea agglomerans. Septic inflammation caused by plant thorn injury can result not only from bacteria. Medical literature indicates that thorns, spines or prickles also introduce pathogenic fungi into animals or humans. Dermatophytes that cause subcutaneous mycoses are unable to penetrate the skin and must be introduced into the subcutaneous tissue by a puncture wound. The common microorganism-thorn combinations seem to have been an important contributor to the fact that so many plant thorns are aposematically colored, as a case of convergent evolution of aposematism in these organisms.Key Words: aposematism, herbivory, pathogen, spine, thorn, bacillus anthracis, clostridium perfringens, sporotrichosis, Mycetoma, subcutaneous mycotic disease     

Macroalgal palatability and the flux of ciguatera toxins through marine food webs

The benthic dinoflagellate Gambierdiscus toxicus produces polyether toxins that cause ciguatera fish poisoning in humans. The toxins initially enter food webs when fish forage on macroalgae, or other substrates, hosting this epiphytic dinoflagellate. Population studies of G. toxicus and risk assessments in ciguatera-prone regions often rely on quantifying dinoflagellates on macroalgae. Underlying these studies is the assumption that the algae sampled represent a readily consumable resource equally available for benthic grazers. However, many algal hosts of G. toxicus possess a variety of defenses against grazing, and host–dinoflagellate associations may act as toxin sources or sinks depending on their palatability. Marine macroalgae may tolerate or avoid herbivory by exhibiting fast growth, by having poor nutritional quality, by utilizing spatial or temporal escapes or by using chemical or structural defenses. Thus, rapidly consumed algae that cope with herbivores by growing fast, such as many filamentous turfs, could be responsible for a high toxin flux even at low dinoflagellate densities. In contrast, ubiquitous unpalatable algae with much higher dinoflagellate densities might contribute little to toxin flux, and effectively act as refuges for G. toxicus. To date, G. toxicus has been reported from 56 algal genera, two cyanobacteria, one diatom, and one seagrass; 63% of these contain species that are defended from fish grazing and other grazers via chemical, morphological or structural defenses, by low nutritional quality, or by a combination of defensive strategies. High dinoflagellate densities on unpalatable macroalgae could indicate passive accumulation of cells on undisturbed hosts, rather than population explosions or active toxin sources for food webs. Understanding the flow of ciguatoxins in nature requires consideration of the ecology of both G. toxicus and its algal hosts. The complexity of marine algal–herbivore interactions also has consequences for other benthic dinoflagellates that produce toxins, which accumulate in consumers.