Dynamics of Ni-based defence and organic defences in the Ni hyperaccumulator, Streptanthus polygaloides (Brassicaceae)

Plants use chemical defences to reduce damage from herbivores and the effectiveness of these defences can be altered by biotic and abiotic factors, such as herbivory and soil resource availability. Streptanthus polygaloides , a nickel (Ni) hyperaccumulator, possesses both Ni-based defences and organic defences (glucosinolates), but the extent to which these defences interact and respond to environmental conditions is unknown. S. polygaloides plants were grown on high-Ni and low-Ni soil and concentrations of Ni and glucosinolates were compared with those of the congeneric non-hyperaccumulator, S. insignus spp. insignus , grown under the same conditions. Ni contents were highest (4000 μg g⁻¹ dry tissue) in S. polygaloides plants grown on high-Ni soil. Glucosinolate content was significantly higher in S. insignus than in S. polygaloides suggesting that plants defended by Ni produce a lower concentration of organic defences. In a separate experiment, high-Ni S. polygaloides plants were exposed to simulated herbivory or live folivores to determine the inducibility of Ni-based and organic defences. Contents of Ni were not affected by either herbivory treatment, whereas glucosinolate concentrations were >30% higher in damaged plants. We concluded that the Ni-based defence of S. polygaloides is not induced by herbivory.     

Variation in Nickel Content in the Nickel-Hyperaccumulating Shrub Psychotria douarrei (Rubiaceae) from New Caledonia1

Plants that hyperaccumulate Ni contain > 1000 ppm (dry wt.) in their tissues. Variation of Ni content within hyperaccumulating plant species is poorly explored. Using the Ni-hyperaccumulating shrub Psychotria douarrei, we documented variation of leaf Ni levels within individual shrubs, and variation with respect to plant size and leaf age. Plant size did not correlate significantly with leaf Ni content, and leaf Ni content did not correlate significantly with soil Ni content. Older leaves contained twice as much Ni as younger leaves. Older leaves also contained greater concentrations of Ca, Fe, and Cr but less K, P, and Cu. Five elements (Zn, Pb, Co, Mn, Mg) showed no significant variation due to leaf age. We also examined the effect of leaf age on epiphyll cover, finding increased epiphyll cover on the upper surface of older leaves. The dominant leafy liverwort epiphyll had a relatively high Ni content (400 ppm), suggesting that epiphylls of Ni hyperaccumulators obtain some Ni from host leaves. Individual shrubs differed in mean leaf Ni content almost two-fold (14,900-27,700 ppm). Variation among branches within individuals also ranged widely; however, this intraplant variability was not strongly correlated with the mean leaf Ni content of an individual shrub. We concluded that Ni contents in leaves of P. douarrei vary considerably due to leaf age, among individual shrubs, and among branches within a shrub.     

Direct and indirect defences induced by piercing-sucking and chewing herbivores in Medicago truncatula

Direct and indirect defences against feeding induced by chewing (Spodoptera littoralis) and piercing-sucking (Tetranychus urticae) herbivores, as well as components of signal transduction, were investigated in the model legume Medicago truncatula. Emitted volatiles, representing a mechanism of indirect defence, were measured and identified by gas chromatography/mass spectrometry (GC-MS). As elements of direct defence, the accumulation of phenolic compounds and of reactive oxygen species (ROS) was assessed using microscopic techniques. Jasmonic acid (JA) and salicylic acid (SA) concentrations were assessed as putative components of signal transduction. Volatile profiles revealed a sizeable number of different substances emitted, particularly sesquiterpenoids. The qualitative composition clearly differed depending on the type of herbivory. The same held true for JA and SA concentrations. Also, deposition of phenolic compounds and the production of ROS around the wounding sites could be detected. Conspicuous differences were found in indirect defence and signalling for different types of herbivory. In contrast, no divergence in direct defences was observed; furthermore, the traits investigated exhibited striking similarities to reactions known to occur upon pathogen attack.     

Herbivory and the evolution of divaricate plants: Structural defences lost on an offshore island

Many island plants are characterized by unique morphology. For example, the high branching angles and small leaves of divaricate plants are a common feature of the New Zealand flora. The divaricate growth form may be an adaptation to deter browsing by extinct avian herbivores (moa); alternatively aspects of the insular climate may be responsible. However, our understanding of the selective pressures responsible for the high branching angles and small leaves of divaricate plants is incomplete. Here, I tested for differences in traits associated with the divaricate growth form between plants from Chatham Island and the New Zealand mainland. Moa never reached the Chatham Islands and its flora is derived from plants on mainland New Zealand. Therefore, I predicted Chatham Island plants to have lost morphological adaptations that may have deterred moa herbivory. Traits were quantified on 316 individuals in the field, allowing for 12 island‐mainland taxonomic comparisons. Chatham Island plants consistently produced smaller branching angles, larger leaves, shorter internodes and larger stems than related mainland plants. Results are therefore consistent with the hypothesis that selection for small leaves and high angled branching may be relaxed on the Chatham Islands due to an absence of moa. Smaller branching angles and larger leaves may offer a competitive advantage to Chatham Island plants.     

Herbivore‐induced plant responses in Brassica oleracea prevail over effects of constitutive resistance and result in enhanced herbivore attack

1. Plant responses to herbivore attack may have community‐wide effects on the composition of the plant‐associated insect community. Thereby, plant responses to an early‐season herbivore may have profound consequences for the amount and type of future attack. 2. Here we studied the effect of early‐season herbivory by caterpillars of Pieris rapae on the composition of the insect herbivore community on domesticated Brassica oleracea plants. We compared the effect of herbivory on two cultivars that differ in the degree of susceptibility to herbivores to analyse whether induced plant responses supersede differences caused by constitutive resistance. 3. Early‐season herbivory affected the herbivore community, having contrasting effects on different herbivore species, while these effects were similar on the two cultivars. Generalist insect herbivores avoided plants that had been induced, whereas these plants were colonised preferentially by specialist herbivores belonging to both leaf‐chewing and sap‐sucking guilds. 4. Our results show that community‐wide effects of early‐season herbivory may prevail over effects of constitutive plant resistance. Induced responses triggered by prior herbivory may lead to an increase in susceptibility to the dominant specialists in the herbivorous insect community. The outcome of the balance between contrasting responses of herbivorous community members to induced plants therefore determines whether induced plant responses result in enhanced plant resistance.     

Testing the optimal defence hypothesis for two indirect defences: extrafloral nectar and volatile organic compounds

Many plants respond to herbivory with an increased production of extrafloral nectar (EFN) and/or volatile organic compounds (VOCs) to attract predatory arthropods as an indirect defensive strategy. In this study, we tested whether these two indirect defences fit the optimal defence hypothesis (ODH), which predicts the within-plant allocation of anti-herbivore defences according to trade-offs between growth and defence. Using jasmonic acid-induced plants of Phaseolus lunatus and Ricinus communis, we tested whether the within-plant distribution pattern of these two indirect defences reflects the fitness value of the respective plant parts. Furthermore, we quantified photosynthetic rates and followed the within-plant transport of assimilates with (13)C labelling experiments. EFN secretion and VOC emission were highest in younger leaves. Moreover, the photosynthetic rate increased with leaf age, and pulse-labelling experiments suggested transport of carbon to younger leaves. Our results demonstrate that the ODH can explain the within-plant allocation pattern of both indirect defences studied.     

Anatomical defences against bark beetles relate to degree of historical exposure between species and are allocated independently of chemical defences within trees

Conifers possess chemical and anatomical defences against tree‐killing bark beetles that feed in their phloem. Resins accumulating at attack sites can delay and entomb beetles while toxins reach lethal levels. Trees with high concentrations of metabolites active against bark beetle‐microbial complexes, and more extensive resin ducts, achieve greater survival. It is unknown if and how conifers integrate chemical and anatomical components of defence or how these capabilities vary with historical exposure. We compared linkages between phloem chemistry and tree ring anatomy of two mountain pine beetle hosts. Lodgepole pine, a mid‐elevation species, has had extensive, continual contact with this herbivore, whereas high‐elevation whitebark pines have historically had intermittent exposure that is increasing with warming climate. Lodgepole pine had more and larger resin ducts. In both species, anatomical defences were positively related to tree growth and nutrients. Within‐tree constitutive and induced concentrations of compounds bioactive against bark beetles and symbionts were largely unrelated to resin duct abundance and size. Fewer anatomical defences in the semi‐naïve compared with the continually exposed host concurs with directional differences in chemical defences. Partially uncoupling chemical and morphological antiherbivore traits may enable trees to confront beetles with more diverse defence permutations that interact to resist attack.     

Conifer responses to a stylet‐feeding invasive herbivore and induction with methyl jasmonate: impact on the expression of induced defences and a native folivore

相似文献   

Leaf trichome responses to herbivory in willows: induction, relaxation and costs

Direct measurement of the carbon (C) 'cost' of mycorrhizas is problematic. Although estimates have been made for arbuscular and ectomycorrhizal symbioses, these are based on incomplete budgets or indirect measurements. Furthermore, the conventional model of unidirectional plant-to-fungus C flux is too simplistic. Net fungus-to-plant C transfer supports seedling establishment in c. 10% of plant species, including most orchids, and bidirectional C flows occur in ectomycorrhiza utilizing soil amino acids. Here, the C cost of mycorrhizas to the green orchid Goodyera repens was determined by measurement of simultaneous bidirectional fluxes of 14C labelled sources using a monoxenic system with the fungus Ceratobasidium cornigerum. Transfer of C from fungus to plant ('up-flow') occurs in the photosynthesizing orchid G. repens (max. 0.06 microg) whereas over five times more current assimilate (min. 0.355 microg) is simultaneously allocated in the reverse direction to the mycorrhizal fungus ('down-flow') after 8 d. Carbon is transferred rapidly, being detected in plant-fungal respiration within 31 h of labelling. This study provides the most complete C budget for an orchid-mycorrhizal symbiosis, and clearly shows net plant-to-fungus C flux. The rapidity of bidirectional C flux is indicative of dynamic transfer at an interfacial apoplast as opposed to reliance on digestion of fungal pelotons.     

Escape behaviour and ultimate causes of specific induced defences in an anuran tadpole

Induced defences, such as the predator avoidance morphologies in amphibians, result from spatial or temporal variability in predation risk. One important component of this variability should be the difference in hunting strategies between predators. However, little is known about how specific and effective induced defences are to different types of predators. We analysed the impact of both pursuing (fish, Gasterosteus aculeatus) and sit-and-wait (dragonfly, Aeshna cyanea) predators on tadpole (Rana dalmatina) morphology and performance (viz locomotive performance and growth rate). We also investigated the potential benefits of the predator-induced phenotype in the presence of fish predators. Both predators induced deeper tail fins in tadpoles exposed to threat of predation, and stickleback presence also induced longer tails and deeper tail muscles. Morphological and behavioural differences resulted in better escape ability of stickleback-induced tadpoles, leading to improved survival in the face of stickleback predation. These results clearly indicate that specific morphological responses to different types of predators have evolved in R. dalmatina. The specific morphologies suggest low correlations between the traits involved in the defence. Independence of traits allows prey species to fine-tune their response according to current predation risk, so that the benefit of the defence can be maximal.     

The spiny,thorny and prickly plants in the flora of Israel

To examine whether spininess evolved at random or differently in various life forms and plant organs, we characterized spiny, thorny and prickly organs in the entire wild flora of Israel (294 such species). Of the species, 63.3% defended their reproductive organs (the most‐defended organ) and 67.0% defended various non‐reproductive organs. Ninety‐three species defended both their reproductive organs and at least one other part; 48.3% defended their leaves and 36.4% their stems and branches. Spiny wings defended stems and branches only in herbaceous (annual or perennial) species. There were clear differences between the life forms. Annuals and perennial herbs defended mostly their reproductive organs (95.7 and 83.0%, respectively), dwarf shrubs defended mostly their leaves (54.2%) and shrubs and trees mostly their branches (89.7 and 76.2%, respectively). Trees do not defend their reproductive organs by associated sharp appendages. The differences in defence on various organs among different life forms may influence the results of meta‐analysis studies of the optimal defence allocation if such differences are not taken into account. We noted spine, thorn and prickle colours for 167 species with yellow, red, orange and white being the dominant, supporting hypotheses about spines being visually aposematic. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 344–352.     

Within and between population variation for zinc and nickel accumulation in two species of Thlaspi (Brassicaceae)

In this study, the differences in zinc (Zn) and nickel (Ni) hyperaccumulation were investigated between three populations of Thlaspi pindicum together with genetic variation within populations of T. pindicum and Thlaspi alpinum var. sylvium, both serpentine endemics. Three experiments were conducted under standard conditions in hydroponic assay. Each experiment contained three treatments of metal: 100 microm Zn, 100 microm Ni, and combined 100/100 microm Zn/Ni. Genetic variation within populations was determined using maternal families. No genetic variation within populations was found for either Zn or Ni hyperaccumulation for both T. pindicum and T. alpinum var. sylvium, but differences were observed for both Zn and Ni hyperaccumulation between populations of T. pindicum. In combined Zn/Ni treatments, Zn inhibited Ni translocation in both species, which is unexpected considering that these species are serpentine endemics and well known Ni hyperaccumulators. The lack of genetic variation for metal hyperaccumulation is possibly due to inbreeding. Since Zn hyperaccumulation is not manifested in the field, inadvertent uptake of Zn is a plausible hypothesis for its preferential uptake.     

Breeding system and floral morphometry of distylous Psychotria L. species in the Atlantic rain forest, SE Brazil

General patterns of floral morphology and incompatibility mechanisms have been described for many distylous plants. The absence of these patterns in typically distylous groups, as observed especially in tropical environments, is interpreted as atypical distyly, or as a new reproductive strategy derived from it. Data are presented here on the morphological and compatibility relations between floral morphs of four Psychotria dimorphic species in the Atlantic rain forest in SE Brazil: Psychotria jasminoides, P. birotula, P. mapourioides, and P. pubigera. When significant differences were found, floral parts were larger in thrum flowers. Results of controlled crosses showed that most incompatible pollen tubes were arrested in the stigma, and only in a low proportion in the upper parts of the style. We conclude that, at the study site, the majority and most important morphological and mating features of typical distyly seem to be conserved in P. jasminoides and P. mapourioides, which presented reciprocal herkogamy, self and intramorph incompatibility, and a balanced morph ratio in the population. Typical distyly in P. birotula is supported by floral morphology, pollen tube data and morph ratio and, in P. pubigera, only by floral morphology and pollen tube data.     

Dioecy derived from distyly and pollination in Psychotria rubra (Rubiaceae) occurring in the Ryukyu Islands,Japan

Floral morphology and breeding system were examined in natural populations of Psychotria rubra (Rubiaceae) occurring in the Ryukyu Islands in southwest Japan. This species is morphologically dimorphic with long‐ and short‐styled morphs; however, these morphs are functionally dioecious. All long‐styled morphs set fruits, but their short anthers completely lack pollen grains; thus, they function as pistillate flowers. Alternately, short‐styled morphs usually have fertile pollen and never set fruit, whether after open or obligate pollinations; thus, they function as staminate flowers. However, some short‐styled morphs in Mounts Katsuu‐dake and Oppa‐dake populations have no fertile pollens. No fruit was obtained from bagging experiments, indicating that P. rubra requires pollinators for its reproduction. In our pollinator observations, flies and short‐tongued wasps were the main visitors of the flowers of P. rubra. Thus, these insects are probably effective pollinators for P. rubra in the Ryukyu Islands. Considering the morphological features, the dioecy found in P. rubra is probably derived from distyly; however, no sufficient evidence is available showing that dioecy has evolved “directly” from distyly.     

Costs and benefits of defences induced by predators differing in dangerousness

While theoretical studies predict that inducible defences should be fine-tuned according to the qualities of the predator, very few studies have investigated how dangerousness of predators, i.e. the rate at which predators kill prey individuals, affects the strength of phenotypic responses and resulting benefits and costs of induced defences. We performed a comprehensive study on fitness consequences of predator-induced responses by involving four predators (leech, water scorpion, dragonfly larva and newt), evaluating costs and benefits of responses, testing differences in dangerousness between predators and measuring responses in several life history traits of prey. We raised Rana dalmatina tadpoles in the presence of free-ranging predators, in the presence of caged predators, and exposed naive and experienced tadpoles to free-ranging predators. Tadpoles adjusted the intensities of their behavioural and morphological defences to predator dangerousness. Survival was lower in the nonlethal presence of the most dangerous predator, while we could not detect costs of induced defences at or after metamorphosis. When exposed to free-ranging predators, small, but not large, tadpoles benefited from exhibiting an induced phenotype in terms of elevated survival when compared to naive tadpoles, but we did not observe higher survival either in tadpoles exhibiting more extreme phenotypes or in tadpoles exposed to the type of predator they were raised with. These results indicate that while predator-induced defences can mirror dangerousness of predators, costs and benefits do not necessarily scale to the magnitude of plastic responses.     

Comparing the qualitatively different effects rapidly evolving and rapidly induced defences have on predator-prey interactions

Interspecific interactions depend not only on the population densities of the interacting species, but on their phenotypes as well. Phenotypic variation can be plastic or heritable and both mechanisms can drive phenotypic change at rates comparable to or faster than those of ecological dynamics (e.g. changes in population abundances or spatial distributions). In this study, we compare the effects rapidly induced and rapidly evolving defences have on community dynamics by considering the fast phenotypic change limit using fast-slow systems theory. Our approach allows us to study phenotypically plastic and evolving systems with one overarching theory, thus capturing the effects rapidly induced defences have on ecological dynamics and how those effects differ from the effects of evolving defences. Our results show that rapidly induced defences tend to stabilize community dynamics and that some behaviours observed in rapidly evolving systems cannot be produced by phenotypic plasticity.     

Pollination and reproduction of Psychotria homalosperma,an endangered distylous tree endemic to the oceanic Bonin (Ogasawara) Islands,Japan

Distyly is usually rare or not observed in species thriving on oceanic islands. The rarity of this breeding system is probably because of the difficulty of colonization for distylous plants and the paucity of pollinators on oceanic islands. However, the endemic and endangered tree Psychotria homalosperma has maintained its distylous nature in the oceanic Bonin Islands, Japan. To understand how the distylous breeding system of P. homalosperma has been maintained on these islands and to characterize the reproduction systems, we studied the pollination and reproductive biology of this species. Specifically, we observed current flower visitors and estimated their effects on plant reproduction. We also examined the floral traits and floral volatiles of P. homalosperma to infer its original pollinators, because plant–pollinator relationships in the Bonin Islands have recently been disrupted by anthropological activities. Finally, we examined the fruit set and pollen tube growth in the stigmas under hand and open pollination. Although original pollinators were presumed to be moths with long proboscises, the introduced honeybee, Apis mellifera, was the most common flower visitor. The honeybee carried pollen grains only unidirectionally, from the short‐ to long‐styled morphs, because it could not reach the hidden stigmas of the short‐styled flowers, and long‐styled flowers set fruits 1.7–38 times more than short‐styled ones. This case study indicates that the instability of pollinator fauna can cause distylous species to be rare on oceanic islands.