Trade-offs between chemical defence and regrowth capacity in Plantago lanceolata

Resistance and tolerance are different strategies of plants to deal with herbivore attack. Since resources are limited and resistance and tolerance serve similar functions for plants, trade-offs between these two strategies have often been postulated. In this study we investigated trade-offs between resistance and one aspect of tolerance, the ability to regrow after defoliation. In order to minimize confounding effects of genetic background and selection history, we used offspring derived from artificial selection lines of ribwort plantain (Plantago lanceolata) that differed in their levels of leaf iridoid glycosides (IGs), allelochemicals that confer resistance to generalist herbivores, to study genetic associations with regrowth ability. We tested whether high-IG plants (1) suffer allocation costs of resistance in terms of reduced shoot and root growth, (2) have reduced regrowth ability (tolerance) after defoliation compared to low-IG plants, and (3) whether such costs are more pronounced under nutrient stress. High-IG plants produced fewer inflorescences and side rosettes than low-IG plants and showed a different biomass allocation pattern, but since neither the vegetative, nor the reproductive biomass differed between the lines, there was no evidence for a cost of IG production in terms of total biomass production under either nutrient condition. High-IG plants also did not suffer a reduced capacity to regrow shoot mass after defoliation. However, after regrowth, root mass of high-IG plants grown under nutrient-poor conditions was significantly lower than that of low-IG plants. This suggests that under these conditions shoot regrowth of high-IG plants comes at a larger expense of root growth than in low-IG plants. We speculate therefore that if there is repeated defoliation, high-IG plants may eventually fail to maintain shoot regrowth capacity and that trade-offs between resistance and tolerance in this system will show up after repeated defoliation events under conditions of low resource availability.     

Grazing tolerance of Gentianella amarella and other monocarpic herbs: why is tolerance highest at low damage levels?

Plants have adapted to compensate for the loss of vegetative biomass and reproductive potential caused by grazing. Shoot damage breaks down the correlative inhibition maintained by apical dominance. The consequent increased branching may lead to increased production of flowers and fruits in damaged plants, provided that enough resources, both in terms of meristems and nutrients, are available. In Gentianella amarella, the removal of the apex of the main stem (10% clipping) had no pronounced effect on branching and plant performance. In one of the two study populations, however, apically damaged plants produced more fruits than undamaged control plants. The plants also fully compensated for 50% removal of the main stem in terms of above-ground biomass, but their fruit production was reduced compared to control and apically damaged plants. After 75% clipping, fruit production was not significantly reduced compared to 50% clipping. Consequently, G. amarella showed highest tolerance in the presence of minor shoot damage. The pattern is qualitatively similar in some other monocarpic species (Gentianella campestris, Erysimum strictum and Rhinanthus minor). Multiple constraints as well as selective forces may shape these compensatory responses: (1) A lack of basal meristems may constrain tolerance of high damage levels. (2) Species with basal meristems may have a potential to tolerate major damage, but a shortage of resources or otherwise unfavourable growth conditions may constrain their compensatory ability. (3) It may be adaptive to have maximum tolerance of low and moderate damage levels if chemical defences reduce the risk of extensive shoot damage as well as the risk of repeated grazing. (4) The compensatory ability of monocarpic species may be affected by selective forces that favour fast vertical growth early in the season and unbranched architecture in undamaged conditions. Therefore, it is not the mere grazing history, but also other factors associated with growth conditions that are required to explain the variation in grazing tolerance.     

No evidence for evolutionarily decreased tolerance and increased fitness in invasive Chromolaena odorata: implications for invasiveness and biological control

Tolerance, the degree to which plant fitness is affected by herbivory, is associated with invasiveness and biological control of introduced plant species. It is important to know the evolutionary changes in tolerance of invasive species after introduction in order to understand the mechanisms of biological invasions and assess the feasibility of biological control. While many studies have explored the evolutionary changes in resistance of invasive species, little has been done to address tolerance. We hypothesized that compared with plants from native populations, plants from invasive populations may increase growth and decrease tolerance to herbivory in response to enemy release in introduced ranges. To test this hypothesis, we compared the differences in growth and tolerance to simulated herbivory between plants from invasive and native populations of Chromolaena odorata, a noxious invader of the tropics and subtropics, at two nutrient levels. Surprisingly, flower number, total biomass (except at high nutrient), and relative increase in height were not significantly different between ranges. Also, plants from invasive populations did not decrease tolerance to herbivory at both nutrient levels. The invader from both ranges compensated fully in reproduction after 50?% of total leaf area had been damaged, and achieved substantial regrowth after complete shoot damage. This strong tolerance to damage was associated with increased resource allocation to reproductive structures and with mobilization of storage reserves in roots. The innately strong tolerance may facilitate invasion success of C. odorata and decrease the efficacy of leaf-feeding biocontrol agents. Our study highlights the need for further research on biogeographical differences in tolerance and their role in the invasiveness of exotic plants and biological control.     

Tolerance to herbivory in lupin genotypes with different alkaloid concentration: Interspecific differences between Lupinus albus L. and L. angustifolius L.

Lupin genotypes accumulate alkaloids that act as feeding deterrents for several kinds of herbivores. Breeding sweet (low alkaloid) genotypes resulted in a greater dependence on pesticides. Besides the concentration of defensive chemicals, plants possess another way to deal with herbivory, to allocate post-damage resources to growth in order to reach compensation in biomass (tolerance).

These two ways to deal with herbivores were postulated as alternative strategies, as scarce resources allocated to one function (growth or secondary metabolism) would not be available for the other function. Genotypes could differ in the way they respond to herbivory; identifying those genotypes with greater ability to overcome the damage would be useful to decrease the use of pesticides.

The aim of this work was to compare tolerance to herbivory in Lupinus albus and Lupinus angustifolius genotypes with contrasting alkaloid concentration. Tolerance was evaluated by comparing growth and grain yield of field-grown cut and uncut plants. Cutting treatments were performed at flowering by cutting 50% of the upper shoot biomass (including the main apex, stems, flowers and leaves). Differences between species were found in their tolerance to herbivory. While L. angustifolius showed full compensation in growth or grain yield that allowed cut plants to equal controls biomass or yield after damage, simulated herbivory reduced growth and grain yield in L. albus.     

Effects of generalist herbivory on resistance and resource allocation by the invasive plant,Phytolacca americana

Successful invasions by exotic plants are often attributed to a loss of co‐evolved specialists and a re‐allocation of resources from defense to growth and reproduction. However, invasive plants are rarely completely released from insect herbivory because they are frequently attacked by generalists in their introduced ranges. The novel generalist community may also affect the invasive plant's defensive strategies and resource allocation. Here, we tested this hypothesis using American pokeweed (Phytolacca americana L.), a species that has become invasive in China, which is native to North America. We examined resistance, tolerance, growth and reproduction of plant populations from both China and the USA when plants were exposed to natural generalist herbivores in China. We found that leaf damage was greater for invasive populations than for native populations, indicating that plants from invasive ranges had lower resistance to herbivory than those from native ranges. A regression of the percentage of leaf damage against mass showed that there was no significant difference in tolerance between invasive and native populations, even though the shoot, root, fruit and total mass were larger for invasive populations than for native populations. These results suggest that generalist herbivores are important drivers mediating the defensive strategies and resource allocation of the invasive American pokeweed.     

How to quantify plant tolerance to loss of biomass?

In some plant species the whole shoot is occasionally removed, as a result of specialist herbivory, grazing, mowing, or other causes. The plant can adapt to defoliation by allocating more to tolerance and less to growth and defense. Plant tolerance to defoliation (TOL1) is typically measured as the ratio between the average dry weight of a group of damaged plants and a control group of undamaged plants, both measured some time after recovery. We develop a model to clarify what TOL1 actually measures. We advocate keeping regrowth (REG2) and shoot–root ratio, both elements of TOL1, separate in the analysis. Based on a resource trade‐off, exotic Jacobaea vulgaris plants from populations in the USA (no specialist herbivory) are expected to grow faster and be less tolerant than native Dutch populations (with specialist herbivory). Indeed Dutch plants had both a significantly larger fraction biomass in roots and faster regrowth (REG2), while US plants attained the highest weight in the control without defoliation. Using key‐factor analysis, we illustrate how growth rates, regrowth, and shoot–root ratio each contribute to final biomass (plant fitness). Our proposed method gives more insight in the mechanisms that underly plant tolerance against defoliation and how tolerance contributes to fitness.     

Evolutionary increases in defense during a biological invasion

Invasive plants generally escape from specialist herbivores of their native ranges but may experience serious damage from generalists. As a result, invasive plants may evolve increased resistance to generalists and tolerance to damage. To test these hypotheses, we carried out a common garden experiment comparing 15 invasive populations with 13 native populations of Chromolaena odorata, including putative source populations identified with molecular methods and binary choice feeding experiments using three generalist herbivores. Plants from invasive populations of C. odorata had both higher resistance to three generalists and higher tolerance to simulated herbivory (shoot removal) than plants from native populations. The higher resistance of plants from invasive populations was associated with higher leaf C content and densities of leaf trichomes and glandular scales, and lower leaf N and water contents. Growth costs were detected for tolerance but not for resistance, and plants from invasive populations of C. odorata showed lower growth costs of tolerance. Our results suggest that invasive plants may evolve to increase both resistance to generalists and tolerance to damage in introduced ranges, especially when the defense traits have low or no fitness costs. Greater defenses in invasive populations may facilitate invasion by C. odorata by reducing generalist impacts and increasing compensatory growth after damage has occurred.     

Infertile times: response to damage in genets of the clonal sedge Carex bigelowii

Tolerance to grazing is a plant trait that can be adaptive in systems where plants are subjected to a diversity of herbivore attack types. To test the tolerance ability of the clonal sedge Carex bigelowii, which is food plant to several herbivores in alpine and arctic areas, and the potential fitness costs of this tolerance, replicated units of genets were subjected to three levels of damage throughout three consecutive seasons. The three levels of treatment were no damage, light damage and heavy damage, and the damage was conducted by tearing off all plant material at 3 and 0 cm above-ground respectively. The genets had no tolerance under damage in terms of sexual reproduction. In terms of clonal reproduction the genets showed tolerance under light damage but not under heavy damage. However, no fitness cost was found for this tolerance ability, i.e. genets had higher reproduction and growth under no damage. The average ramet weight had a similar decrease under both a low and high damage treatment. Changed partitioning of biomass between plant parts and reduced concentration of total non-structural carbohydrates (TNC) in storage organs are possible mechanisms for the ability to uphold clonal reproduction in response to damage. There were no significant indications that tolerance ability or its fitness cost differed between genets. Our results suggest that when subjected to heavy damage genets will only reproduce vegetatively. Consequently, it seems C. bigelowii has evolved to allocate resources to the survival of an already successful genet rather than to a potential new genet of unknown success.     

Local adaptation of annual weed populations to habitats differing in disturbance regime

Plants have evolved several strategies to cope with disturbance, and one strategy is tolerance. In tolerance, plants store resources (meristems, carbohydrates) so that they can resprout after disturbance and thereby compensate to some degree for losses. Because tolerance is costly (it occurs at the expense of current growth), we can expect adaptation to the local disturbance regime. In this study, we determined whether populations of a common European annual weed, Euphorbia peplus, are adapted to the local disturbance regime. We hypothesized that the tolerance and hence compensation for losses in seed and biomass production after experimental damage are greater in plants from more severely disturbed than from less severely disturbed populations. We also hypothesized that transgenerational effects can alter adaptation. We found that compensation for biomass loss to damage was greater for plants from more severely disturbed habitats than for plants from less severely disturbed habitats. This, however, was not at the expense of growth before damage because plants from both disturbance regimes did not show differences when not damaged. Transgenerational effects played a positive role in adaptation to disturbance during germination and maturity. We conclude that local adaptation together with transgenerational effects have evolved in more severely disturbed populations but not in less severely disturbed populations of E. peplus.     

Does cross‐acclimation between drought and freezing stress persist over ecologically relevant time spans? A test using the grass Poa pratensis

• Despite evidence that prior exposure to drought can increase subsequent plant freezing tolerance, few studies have explored such interactions over ecologically relevant time spans. We examined the combined effects of drought and subsequent freezing on tiller growth and leaf sugar concentrations in the grass, Poa pratensis .
• We exposed tillers to no drought (?0.04 MP a), moderate drought (?0.19 MP a) or severe drought (?0.42 MP a) for 3 weeks in summer. Tillers were then frozen in autumn or spring at ?5 °C (frost damage) or at 0 °C (control) for 3 days and harvested after a re‐growth period.
• For shoot growth, there was a significant interaction between drought and autumn freezing, whereby the relative effect of freezing on growth was least for the plants previously exposed to severe drought; however, there was no significant interaction between drought and spring freezing. For root growth, there were no significant interactions between drought and freezing in either season. Leaf sugar concentrations increased significantly with drought intensity, but these effects dissipated within a month, prior to the onset of the autumn freezing treatment.
• Overall, our results suggest that interactions between prior drought and subsequent freezing in P. pratensis may be most relevant in the context of autumn freezing, and despite the important role of soluble sugars in increasing both drought and freezing tolerance in this species, the retention of these compounds after drought stress does not appear to explain the occurrence of drought–frost interactions at ecologically relevant time scales.

     

Overexpression of dehydroascorbate reductase, but not monodehydroascorbate reductase, confers tolerance to aluminum stress in transgenic tobacco

Aluminum (Al) inhibits plant growth partly by causing oxidative damage that is promoted by reactive oxygen species and can be prevented by improving antioxidant capacity. Ascorbic acid (AsA), the most abundant antioxidant in plants, is regenerated by the action of monodehydroascorbate reductase (MDAR) and dehydroascorbate reductase (DHAR). We investigated the role of MDAR and DHAR in AsA regeneration during Al stress using transgenic tobacco (Nicotiana tabacum) plants overexpressing Arabidopsis cytosolic MDAR (MDAR-OX) or DHAR (DHAR-OX). DHAR-OX plants showed better root growth than wild-type (SR-1) plants after exposure to Al for 2 weeks, but MDAR-OX plants did not. There was no difference in Al distribution and accumulation in the root tips among SR-1, DHAR-OX, and MDAR-OX plants after Al treatment for 24 h. However, DHAR-OX plants showed lower hydrogen peroxide content, less lipid peroxidation and lower level of oxidative DNA damage than SR-1 plants, whereas MDAR-OX plants showed the same extent of damage as SR-1 plants. Compared with SR-1 plants, DHAR-OX plants consistently maintained a higher AsA level both with and without Al exposure, while MDAR-OX plants maintained a higher AsA level only without Al exposure. Also, DHAR-OX plants maintained higher APX activity under Al stress. The higher AsA level and APX activity in DHAR-OX plants contributed to their higher antioxidant capacity and higher tolerance to Al stress. These findings show that the overexpression of DHAR, but not of MDAR, confers Al tolerance, and that maintenance of a high AsA level is important to Al tolerance.     

Over-expression of a barley aquaporin increased the shoot/root ratio and raised salt sensitivity in transgenic rice plants

Barley HvPIP2;1 is a plasma membrane aquaporin and its expression was down-regulated after salt stress in barley [Katsuhara et al. (2002) Plant Cell Physiol. 43: 885]. We produced and analyzed transgenic rice plants over-expressing barley HvPIP2;1 in the present study. Over-expression of HvPIP2;1 increased (1) radial hydraulic conductivity of roots (Lp(r)) to 140%, and (2) the mass ratio of shoot to root up to 150%. In these transgenic rice plants under salt stress of 100 mM NaCl, growth reduction was greater than in non-transgenic plants. A decrease in shoot water content (from 79% to 61%) and reduction of root mass or shoot mass (both less than 40% of non-stressed plants) were observed in transgenic plants under salt stress for 2 weeks. These results indicated that over-expression of HvPIP2;1 makes rice plants sensitive to 100 mM NaCl. The possible involvement of aquaporins in salt tolerance is discussed.     

On the correlation between allocation to defence and regrowth in plants

Because storage of resources to regrow after damage and investment in defence draw upon the same resource pool, it has been argued that they should show a negative correlation. We sketch a model for optimal allocation to defence and to storage for regrowth. In the model generalist herbivores exert a constant herbivore pressure against which the plant can defend itself. With discrete intervals, disturbance occurs by an external cause against which the plant cannot defend. This could be an abiotic disturbance or the outbreak of a specialist herbivore that is unaffected by the defence. If we compare genotypes or species, each adapted to its own habitat, then a positive correlation or no correlation between allocation to defence and to regrowth is to be expected. The parameter space in which plants should both defend and store resources for regrowth is limited. Especially under favourable growing conditions, plants should only allocate to growth. We discuss some experimental measures of the regrowth capacity of plants in the context of our model and argue that these should be used with caution.     

Little strokes fell great oaks: minor but chronic herbivory substantially reduces birch growth

Modern concepts of plant tolerance to herbivory are primarily based on studies of short‐term severe damage, whereas the effects of minor chronic damage to long‐lived woody plants, corresponding to background herbivory (2–15% annual loss of foliar biomass in boreal and temperate forests), remain poorly understood. In our experiment, the annual removal of 2, 4, 8 and 16% of the leaf area from naturally growing mountain birch Betula pubescens subsp. czerepanovii saplings during a seven‐year period resulted in a pronounced reduction of plant vertical growth (–30, –34, –45 and –78%, respectively). Leaf size decreased first (already after one year of the 16% treatment), resulting in the reduction of the total leaf area. This effect was followed by a considerable decrease in the length of long shoots in all treatments. Leaf number on the plant was maintained for a longer time, being reduced by the end of the experiment in 16% treatment only; no changes in specific leaf area or chlorophyll fluorescence were observed in either of the treatments. This pattern may indicate that the plant reallocates resources from the growth of the woody parts to the maintenance of the photosynthetic area, and can be seen as a strategy of tolerance to minor herbivory, whereas compensatory responses typical of severe herbivory (increased photosynthesis rates and shoot regrowth) have not been detected. The predicted 2–5% increase in background herbivory due to climate warming can potentially produce previously unrecognised negative impacts on tree growth. We conclude that in the long term, background herbivory is likely to impose stronger effects on the growth of woody plants than short‐term devastating outbreaks of defoliators, thus contributing more to the development of plant evolutionary adaptations to herbivory than severe but episodic bouts of damage.     

葡萄种间杂交砧木育种F_1代植株耐盐性分析

以耐盐性较强的砧木1103P为对照品种,对左山一×SO4杂种砧木F1代的6个株系(A15、A17、A34、A35、A38和A48)及左山一×101-1杂种F1代2个株系(B24和B26)的一年生盆栽扦插苗进行100 mmol·L–1 NaCl胁迫处理,以各自无盐胁迫为对照处理。20天后,根据表型计算盐害指数,测定叶绿素含量、光合气体交换参数、叶绿素荧光参数以及生长量指标;以各项生长指标的耐盐系数为耐盐指标,通过主成分分析、相关性分析、隶属函数分析和聚类分析等方法对葡萄株系进行综合评价。结果表明,A34和A35植株无盐害症状,盐害级数为0;A15和A17植株有少部分叶片边缘焦枯,盐害级数为1。盐胁迫大幅度降低了1103P和B26等株系的叶绿素含量、光合速率、新梢生长量和生物量;而A15、A17、A34和A35植株的各项指标降低幅度较小。将生物量等12个单项指标转换成3个相互独立的综合指标,通过聚类分析,发现A34、A35、A15和A17植株的耐盐性较强,A38、A48和B24植株的耐盐性中等,1103P和B26植株的耐盐性较弱,与盐害分级结果一致。     

Differential invertase activity and root growth between Cu-tolerant and non-tolerant populations in Kummerowia stipulacea under Cu stress and nutrient deficiency

There has been no study on key enzymes in sucrose cleavage in metallophyte plants so far, which may be crucial for the plants’ root growth and heavy-metal tolerance maintenance. Here, we tested the hypothesis that the roots of copper tolerant plants should manifest a higher activity of acid invertases that are rate-limiting in sucrose catabolism than non-tolerant plants both for supporting growth and for their maintaining tolerance under Cu stress. Two populations of Kummerowia stipulacea, one from an ancient waste heap at a Cu mine, and the other from a non-contaminated site, were used in the experiments. The plants were grown in 1/2-fold (control) or 1/20-fold (nutrient deficiency) Hoagland’ solution, with (Cu stress) or without (control) 10 μmol/L Cu²⁺. Plants from the mine proved to be of Cu tolerance. Cu exposure had a stronger inhibition on root growth and thus resulting in a lower root/shoot ratio in the plants of non-mine population compared to the mine population. Cu exposure showed a stronger inhibition of acid invertase activity of Cu non-tolerant plants than Cu-tolerant plants, while neutral/alkaline invertase was insensitive to Cu. A positive correlation between the activity of acid invertases and the root growth and root/shoot ratio was observed. The results indicated an important role of acid invertases in governing root growth and root/shoot biomass allocation in the plants of mine population. The results also suggested that the higher activities in acid invertases of mine population plants might at least partly associate with the plants’ Cu tolerance, and their higher activities in acid invertases in turn played an role in maintenance of the Cu tolerance by supplying carbon and energy for tolerance mechanisms. In addition, the results showed evidence that neutral/alkaline invertase might play a role in compensating for the depression in sucrose catabolism due to Cu-induced inhibition in acid invertases.     

Herbivory resistance traits in populations of Poa ligularis subjected to historically different sheep grazing pressure in Patagonia

Seeding selected populations with high grazing resistance may foster recovery of plant populations threatened by overgrazing. Resistance to grazing depends on grazing avoidance (escape from grazers) and grazing tolerance (ability to growth after defoliation). Many studies of grazing tolerance defoliate plants at a fixed height instead of removing the same proportion of biomass and therefore confound tolerance with avoidance. For this reason, the information on evolution of tolerance to defoliation at the intraspecific level is remarkably scarce despite the abundance of papers published that evaluate responses to defoliation. The estimation of the cost of tolerance is also troublesome because current methods usually include spurious correlations due to correlation between variables that share common terms. The objectives of this paper were to assess the intraspecific variation in tolerance and in traits associated with avoidance and growth in populations with different sheep grazing histories. We also estimated the percentage of biomass removed when the defoliation treatment was imposed at fixed height in order to separate tolerance and avoidance. Finally, we estimated the cost of tolerance using a new method proposed for spurious correlations. Results of a greenhouse experiment indicated no difference in tolerance among the three compared populations. However, the populations from overgrazed fields had more prostrate growth form, higher specific leaf area, and higher tillering rate (when no defoliated) than populations from exclosures. We confirmed that fixed height defoliation would have removed a higher proportion of shoot biomass from taller than from shorter individual plants, confounding grazing tolerance and avoidance. Regarding the cost of tolerance, we found no differences from a null model of no cost, indicating that the evolution (or future breeding) of more tolerant genotypes would not be constrained by this cost.     

Effects of aboveground herbivory on plants with long-term belowground biomass storage

Plant tolerance to herbivory is contingent on multiple traits and adaptive mechanisms, which makes it a complex response with ecological implications. In plants with long-term belowground storage, allocation of biomass to inaccessible parts belowground in response to folivory is a well-recognized tolerance mechanism. In temperate regions, spring growth from buried rootstock is common among winter deciduous plants and is often followed by regrowth after defoliation, both of which draws resources from the stored reserves. We developed a mathematical model to analyze this tolerance response in a winter deciduous plant with long-term belowground biomass when it is defoliated by a specialist insect folivore. The model explores how three closely associated traits—(1) belowground biomass allocation to roots, (2) spring utilization of stored reserves, and (3) post-defoliation regrowth capacity—modulate the persistence and dynamics of the plant and herbivore populations. Model results show that allocation to belowground storage is not only a critical component of tolerance but also influences the herbivore population dynamics in ways that depend on how and when plant biomass is allocated and used. Low belowground biomass allocation and high storage utilization combined with poor photosynthetic growth caused extirpation of the plant population by the defoliating insects. Stable coexistence of the plant at low biomass along with its specialist insect required a moderate amount of post-herbivory belowground allocation. High values of belowground biomass allocation, storage utilization, and photosynthetic growth resulted in sustained cycles of the herbivore and plant populations. Interestingly, utilization of stored reserves had conflicting influence on above and belowground biomass, and strongly affected herbivore population dynamics. Our model thus highlights the complexity of tolerance response when it involves multiple traits and mechanisms as evinced by winter deciduous plants. We close by discussing the implications of our findings for the contributions of defoliating insects to biocontrol programs.     

Protein storage and root:shoot reallocation provide tolerance to damage in a hybrid willow system

To determine the mechanistic basis of tolerance, we evaluated six candidate traits for tolerance to damage using F₂ interspecific hybrids in a willow hybrid system. A distinction was made between reproductive tolerance and biomass tolerance; reproductive tolerance was designated as a plant’s proportional change in catkin production following damage, while biomass tolerance referred to a plant’s proportional change in biomass (i.e., regrowth) following damage. F₂ hybrids were generated to increase variation and independence among candidate traits. Using three clonally identical individuals, pre-damage candidate traits for tolerance to damage (root:shoot ratio, total nonstructural carbohydrate, and total available protein) and post-damage candidate traits (relative root:shoot ratio, phenolic ratio, and specific leaf area ratio) were measured. The range of variation for these six candidate traits was broad. Biomass was significantly increased two years after 50% shoot length removal, and catkin production was not significantly reduced when damaged, suggesting that F₂ hybrids had great biomass tolerance and reproductive tolerance. Based on multiple regression methods, increased reproductive tolerance was associated with increased protein storage and decreased relative root:shoot ratio (reduced root allocation after damage). In addition, a positive relationship between biomass tolerance and condensed tannins was detected, and both traits were associated with increased reproductive tolerance. These four factors explained 57% of the variance in the reproductive tolerance of F₂ hybrids, but biomass tolerance explained the majority of the variance in reproductive tolerance. Changes in plant architecture in response to plant damage may be the underlying mechanism that explains biomass tolerance.     

Branching responses in Silphium integrifolium (Asteraceae) following mechanical or gall damage to apical meristems and neighbor removal

Branching in plants increases plant access to light and provides pathways for regrowth following damage or loss of the apical meristem. We conducted two experiments in an eastern Kansas tallgrass prairie to determine how apical meristem loss (by clipping), apical meristem damage (by insect galling), and increased light availability affected growth, reproduction, and branching in Silphium integrifolium (Asteraceae). The first experiment compared clipping with galling. Clipping increased axillary shoot numbers, while galling increased axillary shoot lengths, reflecting different allocation responses among damage types and inhibition of branching by galls. However, total capitulum production was less in all gall/clip treatments than in intact shoots. The second experiment compared clipping with mowing the surrounding vegetation to increase light availability. Mowing increased total leaf, total capitulum, and axillary shoot length and axillary capitulum production in clipped and unclipped plants and in large vs. small shoots. The presence of the neighboring canopy, not of an intact apical meristem, was therefore the stronger limitation on leaf and capitulum production. These experiments suggest that damage and light competition affected both branching frequency and the partitioning of resources among shoots, branches, and leaves. Because Silphium's growth form is widespread, similar responses may occur in other grassland forbs.