Cascading effects of host size and host plant species on parasitoid resource allocation

1. The bottom‐up factors that determine parasitoid host use are an important area of research in insect ecology. Host size is likely to be a primary cue for foraging parasitoids due to its potential influence on offspring development time, the risk of multiparasitism, and host immunocompetence. Host size is mediated in part by host‐plant traits that influence herbivore growth and potentially affect a herbivore's quality as a host for parasitoids. 2. Here, we tested how caterpillar host size and host plant species influence adult fly parasitoid size and whether host size influences wasp parasitoid sex allocation. We measured the hind tibia lengths and determined the sex of wasp and fly parasitoids reared from 11 common host species of polyphagous caterpillars (Limacodidae) that were in turn reared on foliage of seven different host plant species. 3. We also tested how host caterpillar species, host caterpillar size, and host and parasitoid phenology affect how the parasitoid community partitions host resources. We found evidence that parasitoids primarily partition their shared hosts based on size, but not by host species or phenology. One index of specialisation (d′) supports our observation that these parasitoids are quite generalised within the Limacodidae. In general, wasps were reared from caterpillars collected in early instars, while flies were reared from caterpillars collected in late instars. Furthermore, for at least one species of solitary wasp, host size influenced sex allocation of offspring by ovipositing females. 4. Host‐plant quality indirectly affected the size attained by a tachinid fly parasitoid through its direct effects on the size and performance of the caterpillar host. The host plants that resulted in the highest caterpillar host performance in the absence of enemies also yielded the largest parasitoid flies, which suggests that host plant quality can cascade up to influence the third trophic level.     

Plant allocation of carbon to defense as a function of herbivory,light and nutrient availability

We use modeling to determine the optimal relative plant carbon allocations between foliage, fine roots, anti-herbivore defense, and reproduction to maximize reproductive output. The model treats these plant components and the herbivore compartment as variables. Herbivory is assumed to be purely folivory. Key external factors include nutrient availability, degree of shading, and intensity of herbivory. Three alternative functional responses are used for herbivory, two of which are variations on donor-dependent herbivore (models 1a and 1b) and one of which is a Lotka–Volterra type of interaction (model 2). All three were modified to include the negative effect of chemical defenses on the herbivore. Analysis showed that, for all three models, two stable equilibria could occur, which differs from most common functional responses when no plant defense component is included. Optimal strategies of carbon allocation were defined as the maximum biomass of reproductive propagules produced per unit time, and found to vary with changes in external factors. Increased intensity of herbivory always led to an increase in the fractional allocation of carbon to defense. Decreases in available limiting nutrient generally led to increasing importance of defense. Decreases in available light had little effect on defense but led to increased allocation to foliage. Decreases in limiting nutrient and available light led to decreases in allocation to reproduction in models 1a and 1b but not model 2. Increases in allocation to plant defense were usually accompanied by shifts in carbon allocation away from fine roots, possibly because higher plant defense reduced the loss of nutrients to herbivory.     

Variation in nutrient availability and plant species diversity across forb and graminoid zones of a Northern New England high salt marsh

Plant zonation patterns across New England salt marshes have been investigated for years, but how nutrient availability differs between zones has received little attention. We investigated how N availability, P availability, and plant N status varied across Juncus gerardii, Spartina patens, and mixed forb zones of a Northern New England high salt marsh. We also investigated relationships between several edaphic factors and community production and diversity across the high marsh. P availability, soil salinity, and soil moisture were higher in the mixed forb zone than in the two graminoid zones. NH⁺ ₄-N availability was highest in the J. gerardii zone, but NO^– ₃-N availability and mid season net N mineralization rates did not vary among zones. Plant tissue N concentrations were highest in the mixed forb zone and lowest in the S. patens zone, reflecting plant physiologies more so than soil N availability. Community production was highest in the J. gerardii zone and was positively correlated with N availability and negatively correlated with soil moisture. Plant species diversity was highest in the mixed forb zone and was positively correlated with P availability and soil salinity. Thus, nutrient availability, plant N status, and plant species diversity varied across zones of this high marsh. Further investigation is needed to ascertain if soil nutrient availability influences or is a result of the production and diversity differences that exist between vegetation zones of New England high salt marshes.     

C allocation to the fungus is not a cost to the plant in ectomycorrhizae

Mycorrhizal benefit to plants is most frequently evaluated through growth differences between mycorrhizal (M) and non‐mycorrhizal (NM) plants. These growth differences are often considered to be due to differences in belowground C expenditure, or in cost efficiency, i.e. amount of nutrients acquired per C expended. We searched published reports for relations between plant growth and belowground C allocation, C use efficiency, or nutrient uptake, in ectomycorrhizal (ECM) versus non‐mycorrhizal plants. We found a similar number of cases of negative, null or positive effects of ECM on plant growth. These effects were not correlated with differences on belowground C allocation or C use efficiency between M and NM plants. In contrast, they were very strongly correlated with mycorrhizal effects on plant N gain. A comprehensive analysis of the published data therefore provided evidence that C is an excess, rather than a costly, resource, and that the outcome of the symbiosis depends only on whether mycorrhizae result in increased or decreased nutrient acquisition compared with NM plants, and not on cost efficiency differences between M and NM plants. Consequences of this finding for the regulation of resource exchange between symbionts and the nature of the symbiosis are discussed.     

Sexual allocation in single-flowered hermaphroditic individuals in relation to plant and flower size

Gender expression in hermaphroditic plant species usually departs from strict equisexuality. Study of those departures can aid understanding of non hermaphroditic breeding systems and prevalence of hermaphroditism within angiosperms. Plant size is one of the most studied factors in relation to gender modification. We studied variation in gender expression in the hermaphroditic, mostly single-flowered Paeonia cambessedesii. We separately studied gender modification with increasing plant and flower size using a variety of currencies: number of ovules and stamens, dry mass, N and P. Flower size and number of floral structures (petals, stamens, carpels, and ovules) increased with plant size. Number of ovules increased more rapidly with increasing plant size than number of stamens, indicating a bias towards femaleness with increasing plant size. A similar pattern was found when regressing number of stamens and number of seeds against plant size. Number of floral structures increased with increasing flower mass, but no significant difference was found between stamens and ovules in their rate of increase. Thus, gender modification at plant level was not consistent with patterns at flower level. No differential allocation to stamens vs gynoecium, or sexual structures vs petals was found when using dry mass, N or P as currencies. However, a disproportionate increase in female allocation was found when number of structures was utilised as currency. Study of size-dependent gender expression will benefit from contrast of results obtained using several analysis levels and allocation currencies.     

Diversified impact of mycorrhizal inoculation on mother plants and daughter ramets in the clonally spreading plant Hieracium pilosella L. (Asteraceae)

Plant Ecology - Resource allocation studies of clonal plants whose individuals form networks of interconnected ramets are very challenging. In addition, the presence of mycorrhizal fungi may...     

Effects of intraspecific competition on size variation and reproductive allocation in a clonal plant

Clonal plants grow in diameter rather than height, and therefore competition among genets is likely to be symmetric and to result in smaller variation in size of genets than in non-clonal plants. Moreover, clonal plants can reproduce both sexually and vegetatively. We studied the effects of density on the size of rosettes and of clones, variation in the size of rosettes and of clones, and allocation to sexual and vegetative reproduction in the clonal herb Ranunculus reptans . We grew plants from an artificial population of R. reptans in 32 trays at two densities. After four months, differences in density were still apparent, although clones in the low-density treatment had on average 155% more rosettes and 227% more rooted rosettes than clones in the high-density treatment. The coefficient of variation of these measures of clone size was 15% and 83% higher, respectively, in the low-density treatment. This indicates that intraspecific competition among clones of R. reptans is symmetric and increases the effective population size. Rooted rosettes were larger and varied more in size in the low-density treatment. The relative allocation of the populations to sexual and to vegetative reproduction was 19% and 13% higher, respectively, in the high-density treatment. Moreover, seeds produced in the high-density treatment had a 24% higher mass and a 7% higher germination percentage. This suggests that with increasing density, allocation to sexual reproduction increases more than allocation to vegetative reproduction in R. reptans , which corresponds to the response of some other species with a spreading growth form but not of species with a compact growth form. We conclude that intraspecific competition is an important factor in the life-history evolution of R. reptans because intraspecific competition affects its clonal life-history traits and may affect evolutionary processes such as genetic drift and selection through its effect on the effective population size.     

Estimating plant allometric relationships in a power model with a minimum size of allocation

The allocation of resources among plant structures depends on size. For example, plants need to have a certain minimum size before they allocate resources into producing seeds. Furthermore, the allometric relationship between different plant structures and size has often been found to be adequately described by power functions. Allometric power functions have traditionally led to a bias when estimating and predicting e.g. seed production as a function of size using classical linear statistical methods. The statistical problems of using the linear models when estimating a power function with a threshold value have been solved but due to the relative complexity of the statistical solutions, the solutions are often not used in the ecological literature. Here, an intuitive and simple power model with a minimum size of allocation is investigated using a Bayesian estimation method on a simulated data set. The Bayesian estimation provided satisfactory estimates of the parameters in the model, and the model is suggested as a simple alternative when fitting allometric power functions to ecological data.     

植物组合及水体营养梯度对三种功能性植物生物量累积与分配的影响

设置4个营养水平(I: 0.5 mg N·L^-1, 0.1 mg P·L^-1; II: 1.5 mg N·L^-1, 0.3 mg P·L^-1; III: 4.5 mg N·L^-1, 0.9 mg P·L^-1; Ⅳ: 13.5 mg N·L^-1, 2.7 mg P·L^-1), 研究了水体营养水平、物种组合及其交互作用对入侵漂浮植物凤眼莲、本地扎根浮叶植物黄花水龙和沉水植物苦草生物量累积与分配的影响.结果表明：随营养水平的升高,4个营养水平的凤眼莲和黄花水龙单种和混种的总生物量及茎叶生物量都呈上升趋势,凤眼莲和黄花水龙的总生物量在Ⅲ、Ⅳ处理下平均比Ⅰ、Ⅱ处理下分别增加了54.47%和102.63%;不同植物组合下,苦草各部分生物量呈下降趋势,Ⅲ、Ⅳ处理的总生物量比Ⅰ、Ⅱ处理平均降低了45.88%;经双因素分析,水体营养水平对凤眼莲和黄花水龙生物量有极显著的正影响（P<0.01）,对苦草生物量有极显著的负影响（P<0.01）;而植物组合的影响随目标植物的不 同呈现出差异.     

A simple plant regeneration-ability assay in a range of Lycopersicon species

Plant growth regulator-dependent (PGR-dependent) in vitro shoot organogenesis has been extensively studied in tomato (Lycopersicon esculentum), whereas PGR-independent adventitious shoot organogenesis received marginal attention in L. esculentum and no consideration at all in other Lycopersicon species. In the present study, induction of PGR-independent adventitious shoots was by decapitation of the apex and removal of preexisting shoot meristems of the seedling, and seedling culture on a medium with no PGR supplements. The existence of PGR-independent regeneration-ability was verified in L. esculentum genotypes (high pigment photomorphogenic mutants and wild-type counterparts) and was uncover amongst L. cheesmanii, L. chilense, L. chmielewskii, L. hirsutum, L. parviflorum, L.␣peruvianum and L. pimpinellifolium. Compared to species other than L. esculentum, high pigment photomorphogenic mutants displayed the weakest PGR-independent regeneration-ability. Our results imply that decapitated seedlings cultured on a medium without PGRs can serve as a convenient assay system for genotypic variation in self-controlled, PGR-independent, shoot regeneration-ability in a wide range of Lycopersicon species. Using transverse thin slices of the hypocotyl placed onto a medium supplemented with 0.2 μM zeatin reboside and 0.04 μM IAA, we assessed PGR-mediated shoot regeneration in L. esculentum genotypes. In a given genotype, more plants per seedling were established by PGR-mediated than by PGR-independent regeneration. However, with both modes of organogenesis, only a fraction of shoot buds eventually grew into normal plants, while others developed into abnormal regenerants having no stem. Percentage of stem-deficiency, in a given genotype, was higher in PGR-treated cultures, which indicates that PGRs amplify the formation frequency of imperfect adventitious apical shoot meristems. Unlike L. esculentum, adventitious shoot buds of other Lycopersicon species, induced by wounding seedlings that were not treated with PGRs, rarely formed regenerants lacking a stem.     

Clonal plasticity of aquatic plant species submitted to mechanical stress: escape versus resistance strategy

Background and Aims

The plastic alterations of clonal architecture are likely to have functional consequences, as they affect the spatial distribution of ramets over patchy environments. However, little is known about the effect of mechanical stresses on the clonal growth. The aim of the present study was to investigate the clonal plasticity induced by mechanical stress consisting of continuous water current encountered by aquatic plants. More particularly, the aim was to test the capacity of the plants to escape this stress through clonal plastic responses.

Methods

The transplantation of ramets of the same clone in two contrasting flow velocity conditions was carried out for two species (Potamogeton coloratus and Mentha aquatica) which have contrasting clonal growth forms. Relative allocation to clonal growth, to creeping stems in the clonal biomass, number and total length of creeping stems, spacer length and main creeping stem direction were measured.

Key Results

For P. coloratus, plants exposed to water current displayed increased total length of creeping stems, increased relative allocation to creeping stems within the clonal dry mass and increased spacer length. For M. aquatica, plants exposed to current displayed increased number and total length of creeping stems. Exposure to current induced for both species a significant increase of the proportion of creeping stems in the downstream direction to the detriment of creeping stems perpendicular to flow.

Conclusions

This study demonstrates that mechanical stress from current flow induced plastic variation in clonal traits for both species. The responses of P. coloratus could lead to an escape strategy, with low benefits with respect to sheltering and anchorage. The responses of M. aquatica that may result in a denser canopy and enhancement of anchorage efficiency could lead to a resistance strategy.Key words: Phenotypic plasticity, morphology, submerged aquatic vegetation, clonality, clonal architecture, Potamogeton coloratus, Mentha aquatica, escape, resistance, mechanical stress, thigmomorphogenesis     

Resistance of plant communities to invasion by tall fescue: An experimental study combining species diversity,functional traits and nutrient levels

The grass Festuca arundinacea is often planted for slope stabilisation in South Korea, and is spreading widely beyond the introduction sites. This study used a functional group approach to examine the resistance of plant combinations to invasion by F. arundinacea based on the limiting similarity and diversity-resistance hypotheses, and to elucidate the process of colonisation. The study simulated the environment of construction sites and surrounding areas that might be encountered by expanding populations of F. arundinacea. The role of nutrient condition in the ranking of functional group competitive ability was also examined. Twelve native plant species were categorised into three functional groups using combinations of functional traits. Pairwise (one-to-one competition), multiple (four different neighbouring species) and monoculture experimental settings were designed using two nutrient levels. The Relative Competition Index was used to interpret the competitive effect of neighbouring species on F. arundinacea. Species of the same functional group as F. arundinacea were unable to resist invasion, but annual plants with niche preemption ability could outcompete it. Competitive relationships between native plants and F. arundinacea were explained partially by functional group identity but were inconsistent with the limiting similarity hypothesis and the diversity-interaction. Unforeseen interactions within the artificial communities also produced unexpected effects. In designing artificial plant communities, it is necessary to consider functional traits that reflect the species characteristic of particular periods and indirect effects that modify the interaction between other species.     

Temporal variation of resource allocation between sexual and asexual structures in response to nutrient and water stress in a floating-leaved plant

Aims Adaptive plasticity of biomass allocation to different environmental stressors enables plants to maintain functional relationships among reproductive structures. In freshwater systems, water depth and nutrient content of sediments can have a major effect on biomass allocation in aquatic macrophytes. However, the relative importance of these two stressors is unknown as it is the temporal variation of biomass allocation to the stressors during the growing period. This information may be critical for understanding the tolerance of a plant to environmental conditions.Methods Here, we used four levels of environmental stressors generated by deep and shallow water and high or low sediment nutrient content in a factorial experiment to investigate the biomass allocation responses of a floating macrophyte, Trapella sinensis, during the growing period.Important findings The results showed that the lower sediment nutrient content inhibited biomass increase, whereas the lower water depth increased the sexual reproduction of the plants. The lower sediment content also led to a delay in flowering and compensated trade-offs among sexual reproduction and elongation and clonal reproduction during the growing period. These results indicated that water depth affected the ratio of biomass allocation, whereas the sediment nutrient content affected biomass accumulation when the plants faced these two environmental factors simultaneously. The temporal changes in allocation under lower sediment nutrient content underscored the importance of collecting data at different stages of growth when trying to interpret resource allocation, especially in resource-limited environments.