Abiotic stress mediates top-down and bottom-up control in a Southwestern Atlantic salt marsh

Increasing evidence has shown that nutrients and consumers interact to control primary productivity in natural systems, but how abiotic stress affects this interaction is unclear. Moreover, while herbivores can strongly impact zonation patterns in a variety of systems, there are few examples of this in salt marshes. We evaluated the effect of nutrients and herbivores on the productivity and distribution of the cordgrass Spartina densiflora along an intertidal stress gradient, in a Southwestern Atlantic salt marsh. We characterized abiotic stresses (salinity, ammonium concentration, and anoxia) and manipulated nutrients and the presence of the herbivorous crab Neohelice (Chasmagnathus) granulata, at different tidal heights with a factorial experiment. Abiotic stress increased at both ends of the tidal gradient. Salinity and anoxia were highest at the upper and lower edge of the intertidal, respectively. Nutrients and herbivory interacted to control cordgrass biomass, but their relative importance varied with environmental context. Herbivory increased at lower tidal heights to the point that cordgrass transplants onto bare mud substrate were entirely consumed unless crabs were excluded, while nutrients were most important where abiotic stress was reduced. Our results show how the impact of herbivores and nutrients on plant productivity can be dependent on environmental conditions and that the lower intertidal limits of marsh plants can be controlled by herbivory.     

Relative Importance of Environmental Stress and Herbivory in Reducing Litter Fall in a Semiarid Woodland

We examined the impact of soil stress (low water and nutrient availabilities) and two keystone insect herbivores on pinyon pine (Pinus edulis) needle litterfall. We compared trees growing on two distinct soil types: volcanic cinders, which exhibit pronounced water and nutrient limitation, and sandy-loam soils, which have higher water-storage capacity and nutrient availability. Using two long-term herbivore removal experiments (15 and 18 years, respectively), we also examined the effects of the pinyon needle scale (Matsucoccus acalyptus, which attacks juvenile trees) and the stem-boring moth (Dioryctria albovittella, which attacks mature trees) on pinyon litterfall. These herbivores reach high densities on cinder soils but are absent or occur at much lower levels on sandy-loam soils. Four years of litterfall measurements showed four major patterns. First, independent of herbivory, needle litterfall was 20% lower under trees on high-stress cinder soils than on sandy-loam soils. Second, in agreement with the negative impact of scales on tree growth (that is, a 30% decline in stem growth), trees with scale infestations had 25% lower litterfall rates than trees resistant to scale; however, 15 years of scale-insect removal did not significantly increase needle litterfall. This implies possible intrinsic differences in litter production between scale-resistant and scale-susceptible trees. Third, in contrast with significant negative effects of moth herbivory on tree growth (that is, a 27% decline in stem growth), moth herbivory had no effect on needle litterfall. This, along with increased stem density in moth-susceptible trees, may be evidence of compensatory production. Fourth, there were strong year by soil type and year by scale herbivory interactions, such that in some years the effect on litterfall can be obscured or reversed by some other factor. In summary, soil stress has a strong and predictable effect on needle litterfall, whereas the relationship between insect herbivory and needle litterfall is weaker and depends on the individual herbivore. These effects, however, are mediated by other environmental factors that have considerable annual variation.     

Crab herbivory regulates re‐colonization of disturbed patches in a southwestern Atlantic salt marsh

Recent work exploring the effects of physical stress and herbivory on secondary succession in estuarine plant communities agrees with basic stress models and reveal that herbivory is an important force in brackish and oligohaline marshes but negligible in physically stressful salt marshes. In these systems, herbivores are terrestrial, and thus negatively affected by the same stressful factors that affect marsh plants (i.e. frequent flooding or high salinities). We evaluated the effects of a marine herbivore (i.e. the crab Neohelice granulata) on plant secondary succession in a southwestern Atlantic salt marsh. Field surveys revealed that disturbance‐generated bare patches have harsh physical conditions and that their edges suffer higher herbivore pressure compared to the marsh matrix. A factorial experiment demonstrated that asexual expansion of the surrounding plants is the only possible mechanism to re‐colonize disturbed patches and that crab exclusion can increase this colonization rate by more than 30 times. Our results show that even in highly stressful environments, herbivores strongly impact marsh structure by regulating patch recovery. The synergism of physical stress and herbivory may make plant succession an extremely slow process and lead to the prevalence of bare areas.     

Effects of flooding, salinity and herbivory on coastal plant communities, Louisiana, United States

Flooding and salinity stress are predicted to increase in coastal Louisiana as relative sea level rise (RSLR) continues in the Gulf of Mexico region. Although wetland plant species are adapted to these stressors, questions persist as to how marshes may respond to changed abiotic variables caused by RSLR, and how herbivory by native and non-native mammals may affect this response. The effects of altered flooding and salinity on coastal marsh communities were examined in two field experiments that simultaneously manipulated herbivore pressure. Marsh sods subjected to increased or decreased flooding (by lowering or raising sods, respectively), and increased or decreased salinity (by reciprocally transplanting sods between a brackish and fresh marsh), were monitored inside and outside mammalian herbivore exclosures for three growing seasons. Increased flooding stress reduced species numbers and biomass; alleviating flooding stress did not significantly alter species numbers while community biomass increased. Increased salinity reduced species numbers and biomass, more so if herbivores were present. Decreasing salinity had an unexpected effect: herbivores selectively consumed plants transplanted from the higher-salinity site. In plots protected from herbivory, decreased salinity had little effect on species numbers or biomass, but community composition changed. Overall, herbivore pressure further reduced species richness and biomass under conditions of increased flooding and increased salinity, supporting other findings that coastal marsh species can tolerate increasingly stressful conditions unless another factor, e.g., herbivory, is also present. Also, species dropped out of more stressful treatments much faster than they were added when stresses were alleviated, likely due to restrictions on dispersal. The rate at which plant communities will shift as a result of changed abiotic variables will determine if marshes remain viable when subjected to RSLR. Received: 8 April 1998 / Accepted: 15 June 1998     

Tracking sucking herbivory with nitrogen isotope labelling: Lessons from an individual trait-based approach

Response and effect traits help to understand how changes in ecological communities (e.g. in response to land use) relate to changes in ecosystem functioning. In grasslands, plants and insect herbivores are involved in many ecosystem processes such as herbivory and plant biomass production. Simultaneous changes in the trait composition of both plants and herbivores should affect herbivory rates, with consequences for plant growth and potentially biomass production. The mechanisms underlying these links are little understood for grasses and sucking insects, which build a major part of grassland communities. In a mesocosm experiment, we manipulated the composition of grasses and sucking herbivores (Hemiptera) to study the role of plant traits, herbivore traits and their interaction on herbivory and plant growth. Because sucking herbivory is generally difficult to quantify, we developed a novel experimental setting, in which we labelled plants with ¹⁵N isotope. This allowed to quantify ¹⁵N uptake and thus sucking rates of individuals. We found that herbivory and simultaneous plant growth reduction are most strongly linked to herbivore species identity. Unexpectedly, herbivory did not increase with herbivore size, but was highest for small species and for thin-bodied Heteroptera. Additionally, herbivory and plant growth reduction depended on the interacting herbivore and plant species, indicating trait matching, which could, however, not be explained with commonly used traits. This indicates that mechanisms linking ecological communities and ecosystem processes are highly context-specific. To understand how global change affects ecosystem functioning, studies need to cover all functionally relevant groups, including plant sap suckers.     

Herbivory-induced overcompensation and resource-dependent production of extrafloral nectaries in Luffa cylindrica (Cucurbitaceae)

Extrafloral nectaries (EFNs) are nectar secretory structures involved in the indirect defense of plants. In the sponge gourd (Luffa cylindrica), EFNs commonly occur on the lower surface of leaf blades and stipules and remain functional until leaf senescence. To test the hypothesis that the development of EFNs is influenced by herbivore damage and resource availability, we grew Luffa cylindrica under different concentrations of Hoagland's nutrient solution (nutrient-poor conditions: 10%, 50%; and control condition: 100%) and two herbivory treatments (damaged and undamaged leaves). We collected ten leaves from treated plants to quantify leaf area and EFN density. Overall, leaf area increased and EFN decreased in damaged plants, but this significantly depended on nutritional status. In undamaged plants, EFN density tended to remain constant, whereas foliar area increased with nutrient input. Under herbivory, foliar area increased at 10% but decreased at 50 and 100% of nutrients in relation to undamaged plants, whereas EFN density tended to increase with nutrient availability to exceed undamaged plants under control concentrations. Plants under nutrient-poor conditions subjected to herbivory exhibited an increased foliar area, characterizing a compensatory mechanism. Our results suggest that herbivore-induced indirect defense is a damage- and resource-dependent response in Luffa cylindrica. These findings contribute to understanding the factors that modulate indirect defenses and plant-herbivore-environment interactions.     

Upscaling hypotheses on herbivore damage in plants facing environmental stress: Variation among scales and plant enemies in a relict tree

The few existing analyses of broad-scale variation in plant–herbivore interactions are largely exploratory and not based on hypotheses on the effect of varying environmental conditions on the patterns of herbivore damage. However, there are two established hypotheses that relate herbivore damage and environmental stress mostly tested at the within-population level: the Plant Stress Hypothesis (PSH) and the Plant Vigor Hypothesis (PVH). To examine if support for the PSH holds true at broader scales, we tested it at the micro- (among individuals), meso- (between habitats) and macroscale (between geographical regions, Iberia and Macaronesia). We measured plant stress and leaf damage by different natural enemies (chewers, miners and pathogens) in 12 populations across the range of the relict tree Prunus lusitanica. Results regarding the proportion of leaves attacked by any enemy showed no support for the PSH at either micro or mesoscale, observing instead higher damage in less stressed individuals within populations and in high-quality habitats. However, the latter was true exclusively within the Macaronesian region, highlighting the important effect of the type of abiotic stress on plant susceptibility to herbivory. At the macroscale, alternatively, results showed support for the PSH: we observed a higher proportion of leaves attacked in Macaronesia than in Iberia, where plants unexpectedly exhibited greater stress symptoms. Among-scale inconsistencies may respond to the effect of distinct climatic factors governing both plant and herbivore dynamics at large scales, particularly in species with distributional ranges across contrasting climatic regions such as P. lusitanica. Finally, different plant enemies showed different patterns of damage, corroborating the need to study them separately to achieve consistent generalizations of these hypotheses.     

Variation in insect herbivory across a salt marsh tidal gradient influences plant survival and distribution

Herbivore damage and impact on plants often varies spatially across environmental gradients. Although such variation has been hypothesized to influence plant distribution, few quantitative evaluations exist. In this study I evaluated patterns of insect herbivory on an annual forb, Atriplex patula var. hastata, across a salt marsh tidal gradient, and performed experiments to examine potential causes and consequences of variation in herbivory. Damage to plants was generally twice as great at mid-tidal elevations, which are more frequently inundated, than at higher, less stressful, elevations at five of six surveyed sites. Field herbivore assays and herbivore preference experiments eliminated the hypothesis that plant damage was mediated by herbivore response to differences in host plants across the gradient. Alternately, greater herbivore densities in the mid-marsh, where densities of an alternate host plant (Salicornia europaea) were high, were associated with greater levels of herbivory on Atriplex, suggesting spillover effects. The effect of insect herbivores on host plant performance varied between the two sites studied more intensively. Where overall herbivore damage to plants was low, herbivory had no detectable effect on plant survival or seed production, and plant performance did not significantly differ between zones. However, where herbivore damage was high, herbivores dramatically reduced both plant survival (>50%) and fruit production (40-70%), and their effects were stronger in the harsher mid-marsh than the high marsh. Thus herbivores likely play a role in maintaining lower Atriplex densities in mid-marsh. Overall, these results suggest that variation in herbivore pressure can be an important determinant of patterns of plant abundance across environmental gradients.     

Interactive effects of salinity and water depth on the growth of Melaleuca ericifolia Sm. (Swamp paperbark) seedlings

Melaleuca ericifolia Sm. (Swamp paperbark) is a common tree species in freshwater and brackish wetlands in southern and eastern Australia. The survival of this species in many wetlands is now threatened by increased salinity and inappropriate water regimes. We examined the response of 5-month-old M. ericifolia seedlings to three water depths (exposed, waterlogged and submerged) at three salinities (2, 49 and 60 dS m⁻¹). Increasing water depth at the lowest salinity did not affect survival, but strongly inhibited seedling growth. Total biomass, leaf area and maximum root length were highest in exposed plants, intermediate in waterlogged plants and lowest in submerged plants. Although completely submerged plants survived for 10 weeks at the lowest salinity, they demonstrated negative growth rates and were unable to extend their shoots above the water surface. At the higher salinities, M. ericifolia seedlings were intolerant of waterlogging and submergence: all plants died after 9 weeks at 60 dS m⁻¹. Soil salinities increased over time, and by Week 10, exceeded external water column salinities in both the exposed and waterlogged treatments. In exposed sediment, ∼90% of plants survived for 10 weeks at 60 dS m⁻¹ even though soil salinities reached ∼76 dS m⁻¹. No mortality occurred in the exposed plants at 49 dS m⁻¹, and small but positive relative growth rates were recorded at Week 10. We conclude that at low salinities M. ericifolia seedlings are highly tolerant of sediment waterlogging, but are unlikely to tolerate prolonged submergence. However, at the higher salinities, M. ericifolia seedlings are intolerant of waterlogging and submergence and died rapidly after 5 weeks exposure to this combination of environmental stressors. This research demonstrates that salinity may restrict the range of water regimes tolerated by aquatic plants.     

Induction of extrafloral nectar depends on herbivore type in invasive and native Chinese tallow seedlings

Although induced defenses are widespread in nature, and a potentially important strategy used by invasive plants, it is unclear how induced defenses vary among populations and whether the intensity and duration of induced defenses depends on herbivore type. For invasive plants, low herbivore loads in their introduced ranges can lead to differences in herbivore defense compared to their native ranges, but we currently know little about how induced defenses vary among native and invasive populations. We conducted a greenhouse experiment to examine variation in one type of induced defense, extrafloral nectar (EFN) production, among native and invasive populations of Chinese tallow tree, Triadica sebifera. We experimentally manipulated herbivory from an exotic generalist scale insect, a native generalist caterpillar, both herbivores, or neither and then examined EFN production by Triadica. Damage from leaf-chewing caterpillars resulted in strongly induced EFN in both native and invasive populations while damage from phloem-feeding scales did not. Extrafloral nectar production and dissolved solute content peaked 4 days after caterpillar herbivory for both native and invasive populations. Number and proportion of leaves producing EFN, EFN volume and concentration of dissolved solutes were similar among native and invasive populations. These results suggest that selection for indirect defenses may be different than selection for other defenses in the introduced ranges of invasive plants, as constitutive and induced EFN production is retained in invasive populations.     

Can plant–natural enemy communication withstand disruption by biotic and abiotic factors?

相似文献   

Nutritional Status as the Key Modulator of Antioxidant Responses Induced by High Environmental Ammonia and Salinity Stress in European Sea Bass (Dicentrarchus labrax)

Salinity fluctuation is one of the main factors affecting the overall fitness of marine fish. In addition, water borne ammonia may occur simultaneously with salinity stress. Additionally, under such stressful circumstances, fish may encounter food deprivation. The physiological and ion-osmo regulatory adaptive capacities to cope with all these stressors alone or in combination are extensively addressed in fish. To date, studies revealing the modulation of antioxidant potential as compensatory response to multiple stressors are rather lacking. Therefore, the present work evaluated the individual and combined effects of salinity challenge, ammonia toxicity and nutritional status on oxidative stress and antioxidant status in a marine teleost, European sea bass (Dicentrarchus labrax). Fish were acclimated to normal seawater (32 ppt), to brackish water (20 ppt and 10 ppt) and to hypo-saline water (2.5 ppt). Following acclimation to different salinities for two weeks, fish were exposed to high environmental ammonia (HEA, 20 mg/L representing 50% of 96h LC₅₀ value for ammonia) for 12 h, 48 h, 84 h and 180 h, and were either fed (2% body weight) or fasted (unfed for 7 days prior to HEA exposure). Results show that in response to decreasing salinities, oxidative stress indices such as xanthine oxidase activity, levels of hydrogen peroxide (H₂O₂) and lipid peroxidation (malondialdehyde, MDA) increased in the hepatic tissue of fasted fish but remained unaffected in fed fish. HEA exposure at normal salinity (32 ppt) and at reduced salinities (20 ppt and 10 ppt) increased ammonia accumulation significantly (84 h–180 h) in both feeding regimes which was associated with an increment of H₂O₂ and MDA contents. Unlike in fasted fish, H₂O₂ and MDA levels in fed fish were restored to control levels (84 h–180 h); with a concomitant increase in superoxide dismutase (SOD), catalase (CAT), components of the glutathione redox cycle (reduced glutathione, glutathione peroxidase and glutathione reductase), ascorbate peroxidase (APX) activity and reduced ascorbate (ASC) content. On the contrary, fasted fish could not activate many of these protective systems and rely mainly on CAT and ASC dependent pathways as antioxidative sentinels. The present findings exemplify that in fed fish single factors and a combination of HEA exposure and reduced seawater salinities (upto 10 ppt) were insufficient to cause oxidative damage due to the highly competent antioxidant system compared to fasted fish. However, the impact of HEA exposure at a hypo-saline environment (2.5 ppt) also defied antioxidant defence system in fed fish, suggesting this combined factor is beyond the tolerance range for both feeding groups. Overall, our results indicate that the oxidative stress mediated by the experimental conditions were exacerbated during starvation, and also suggest that feed deprivation particularly at reduced seawater salinities can instigate fish more susceptible to ammonia toxicity.     

Photophysiological responses of Halophila johnsonii to experimental hyposaline and hyper-CDOM conditions

The endemic seagrass Halophila johnsonii grows intertidally to 3 m deep, in both marine and riverine influenced habitats of eastern Florida. Salinity and chromophoric dissolved organic matter (CDOM) levels widely fluctuate across this broad habitat range, changing tidally and with variable influx of freshwater from watershed runoff, river discharge and stochastic storm events. CDOM exponentially absorbs light in the UV to blue wavelengths, affecting optical water quality. H. johnsonii produces 15 flavonoid compounds that maximally absorb in the UV range. These flavonoids are thought to function as UV-protectants (UVP) in high-light and UV-intense environments. This mesocosm study examined the photosynthetic capacity, quantum efficiency and pigment content of H. johnsonii under experimental treatments of three salinities (10, 20 and 30) with and without CDOM. Main treatment effects and possible interactive effects at both short- (1 day to 1 week) and longer-term (1 month) time scales were examined. There were no significant CDOM or CDOM x salinity effects over any of the experimental treatment durations. There was 100% mortality of plants at salinity 10 after 10 days regardless of water color. UVP content of leaves was not affected by CDOM in this study, but there was significant variation in UVP in response to salinity. Our results do not support the primary role of UVP in this species as a sunscreen, but indicate that different salinity environments contribute to changes in the levels of these flavonoids. The UVP response to salinity stress response was not mitigated by a decrease in UV-radiation (increased CDOM) as H. johnsonii continued to put energy into the production of the carbon-rich flavonoids regardless of potential UV-stress. The experimental results indicate that prolonged hypo-salinity conditions are an important environmental factor to manage in the limited geographic range of H. johnsonii.     

Similar responses of insect herbivores to leaf fluctuating asymmetry

Fluctuating asymmetry (FA) represents small, random variation from symmetry and it has been used as an indicator of plant quality and susceptibility to herbivory. In this study, the effects of FA on the responses of distinct herbivore species belonging to several guilds were examined along an environmental gradient in south Florida. This approach was chosen because it relies on a multi-species approach to the study of fluctuating asymmetry and patterns of herbivory between and within plants along an environmental gradient of salinity and plant stress. To examine differences in FA between and within plant communities, seven plant species were investigated. Four of these plants were coastal species and three species occurred in upland communities. Levels of FA were assessed before herbivory and plants were followed for the whole herbivory season in 2006. Coastal plants exhibited significantly higher salt concentration, higher percentage of asymmetric leaves and higher asymmetry levels than upland plants. Herbivore abundance varied widely amongst the seven species studied, but quantitative syntheses of our results indicated significant and positive responses of insect herbivores to leaf asymmetry: insects were 25.11% more abundant on more asymmetric plants and stronger effects of asymmetry were observed for leaf miners compared to gall-formers. As demonstrated by other recent studies, FA might be used as a reliable stress indicator, leading to similar responses of insect herbivores to variation in leaf symmetry.     

Effects of leaf and root herbivory by potential insect biological control agents on the performance of invasive Vincetoxicum spp.

The European leaf-feeding moth Abrostola asclepiadis and root-feeding beetle Eumolpus asclepiadeus are promising biological control agents for two European swallow-worts (Vincetoxicum rossicum and Vincetoxicum nigrum) in North America, however, their impact on plant performance is uncertain. Densities of each herbivore were manipulated in a common garden to determine whether leaf and root herbivory affect the performance of these plants. During the second year of the experiment, V. rossicum and V. nigrum unexpectedly became infected with the fungal pathogens Ascochyta sp. and Cercospora sp. (Ascomycota), respectively. Although pathogen infection mainly reduced shoot height and delayed reproduction, herbivore effects on plant growth were still evident. Leaf herbivory by A. asclepiadis had no effect on plant growth 1 year after defoliation. Root herbivory by E. asclepiadeus reduced shoot height and plant biomass and decreased the ability of plants to compensate for pathogen attack. Pathogen infection prevented detection of herbivore effect on reproduction. Due to its substantial impact on plant biomass, E. asclepiadeus should be further evaluated as a biological control agent against Vincetoxicum spp. populations invading open habitats in North America. Further research is needed to evaluate the impact of A. asclepiadis in combination with E. asclepiadeus and plant competition under high and low light conditions.     

How do two specialist butterflies determine growth and biomass of a shared host plant?

Although insect herbivory can modify subsequent quantity and quality of their host plants, change in plant quantity following herbivory has received less attention than plant quality. In particular, little is known about how previous herbivore damage determines plant growth and biomass in an insect species-specific manner. We explored whether herbivore species-specific food demand influences plant growth and biomass. To do this, we conducted a series of experiments and field survey using two specialist butterflies, Sericinus montela and Atrophaneura alcinous, and their host plant, Aristolochia debilis. It is known that A. alcinous larva requires four times more food resources to fulfill its development than S. montela larva. Despite that A. alcinous larvae imposed greater damage on plants than S. montela larvae, plant growth did not differ due to herbivory by these species both in single and multiple herbivory events. On the other hand, total aboveground biomass of the plants was reduced more by A. alcinous than S. montela feeding regardless of the number of herbivory events. Feeding on plants with a history of previous herbivory neither decreased nor increased larval growth. Our results suggest that food demand of the two butterfly species determined subsequent plant biomass, although the plant response may depend on tolerance of the host plant (i.e., ability to compensate for herbivore damage). Such difference in the effects of different herbivore species on host plant biomass is more likely to occur than previously thought, because food demand differs in most herbivore species sharing a host plant.     

Experimental Infection of Plants with an Herbivore-Associated Bacterial Endosymbiont Influences Herbivore Host Selection Behavior

Although bacterial endosymbioses are common among phloeophagous herbivores, little is known regarding the effects of symbionts on herbivore host selection and population dynamics. We tested the hypothesis that plant selection and reproductive performance by a phloem-feeding herbivore (potato psyllid, Bactericera cockerelli) is mediated by infection of plants with a bacterial endosymbiont. We controlled for the effects of herbivory and endosymbiont infection by exposing potato plants (Solanum tuberosum) to psyllids infected with “Candidatus Liberibacter solanacearum” or to uninfected psyllids. We used these treatments as a basis to experimentally test plant volatile emissions, herbivore settling and oviposition preferences, and herbivore population growth. Three important findings emerged: (1) plant volatile profiles differed with respect to both herbivory and herbivory plus endosymbiont infection when compared to undamaged control plants; (2) herbivores initially settled on plants exposed to endosymbiont-infected psyllids but later defected and oviposited primarily on plants exposed only to uninfected psyllids; and (3) plant infection status had little effect on herbivore reproduction, though plant flowering was associated with a 39% reduction in herbivore density on average. Our experiments support the hypothesis that plant infection with endosymbionts alters plant volatile profiles, and infected plants initially recruited herbivores but later repelled them. Also, our findings suggest that the endosymbiont may not place negative selection pressure on its host herbivore in this system, but plant flowering phenology appears correlated with psyllid population performance.     

Physiological and growth responses of Centaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability

Summary Centaurea maculosa seedlings were grown in pots to study the effects of root herbivory by Agapeta zoegana L. (Lep.: Cochylidae) and Cyphocleonus achates Fahr. (Col.: Curculionidae), grass competition and nitrogen shortage (each present or absent), using a full factorial design. The aims of the study were to analyse the impact of root herbivory on plant growth, resource allocation and physiological processes, and to test if these plant responses to herbivory were influenced by plant competition and nitrogen availability. The two root herbivores differed markedly in their impact on plant growth. While feeding by the moth A. zoegana in the root cortex had no effect on shoot and root mass, feeding by the weevil C. achates in the central vascular tissue greatly reduced shoot mass, but not root mass, leading to a reduced shoot/root ratio. The absence of significant effects of the two herbivores on root biomass, despite considerable consumption, indicates that compensatory root growth occurred. Competition with grass affected plant growth more than herbivory and nutrient status, resulting in reduced shoot and root growth, and number of leaves. Nitrogen shortage did not affect plant growth directly but greatly influenced the compensatory capacity of Centaurea maculosa to root herbivory. Under high nitrogen conditions, shoot biomass of plants infested by the weevil was reduced by 30% compared with uninfested plants. However, under poor nitrogen conditions a 63% reduction was observed compared with corresponding controls. Root herbivory was the most important stress factor affecting plant physiology. Besides a relative increase in biomass allocation to the roots, infested plants also showed a significant increase in nitrogen concentration in the roots and a concomitant reduction in leaf nitrogen concentration, reflecting a redirection of the nitrogen to the stronger sink. The level of fructans was greatly reduced in the roots after herbivore feeding. This is thought to be a consequence of their mobilisation to support compensatory root growth. A preliminary model linking the effects of these root herbivores to the physiological processes of C. maculosa is presented.     

Effects of short- and long-term salinity on leaf water relations, gas exchange, and growth in Ipomoea pes-caprae

Ipomoea pes-caprae is widespread in pantropical coastal areas along the beach. The aim of this study was to investigate the salinity tolerance level and physiological mechanisms that allow I. pes-caprae to endure abrupt increases in salinity under brief or prolonged exposure to salinity variations. Xylem sap osmolality (X_osm), leaf water relations, gas exchange, and number of produced and dead leaves were measured at short- (1-7 d) and long- (22-46 d) term after a sudden increase in soil salinity of 0, 85, 170, and 255 mM NaCl. In the short-term, X_osm was not affected by salinity, but in the long-term there was a significant increase in plants grown in presence of salt compared with control plants. After salt addition, the plants showed osmotic stress with temporal cell turgor loss. However, the water potential gradient for water uptake was re-established at 4, 7 and 22 d after salt addition, at 85, 170 and 255 mM NaCl, respectively. In the short-term I. pes-caprae was able to tolerate salinities of up to 255 mM NaCl without significant reduction in carbon assimilation or growth. With the duration of stress, leaf ion concentration continued to increase and reached toxic levels at high salinity with a progressive decrease in photosynthetic rate, reduced leaf formation and accelerated senescence. Then, if high levels of soil salts from tidal inundation occur for short periods, the survival of I. pes-caprae is possible, but prolonged exposure to salinity may induce metabolic damage and reduce drastically the plant growth.