Compensatory Regrowth Research Articles

Perfect Compensation Is Sufficient to Explain Insect Outbreaks Previously Attributed to Overcompensation : (A Comment on Stieha et al., "The Effects of Plant Compensatory Regrowth and Induced Resistance on Herbivore Population Dynamics").

AbstractIn "The Effects of Plant Compensatory Regrowth and Induced Resistance on Herbivore Population Dynamics," which appeared in The American Naturalist in 2016, Stieha et al. argued that overcompensatory regrowth of plant tissues lost to herbivory ("overcompensation") promotes cyclic herbivore outbreaks. In contrast, they concluded that partial regrowth ("tolerance") stabilizes herbivore dynamics, preventing outbreaks. These conclusions were based on a comparison between two plant-herbivore models that differed in two properties: (1) whether biomass could ever be higher after herbivory and regrowth than before herbivory (i.e., is overcompensatory regrowth possible?) and (2) how much herbivory the plants could withstand before only being able to partially compensate for losses (for overcompensating plants, there was a threshold herbivory level above which this occurred, whereas tolerant plants always showed partial compensation). While Stieha et al. supposed that difference 1 was responsible for the increased propensity for outbreaks in their overcompensation model, we show here that, in fact, difference 2 is responsible. Thus, we conclude that Stieha et al.'s results about "overcompensating" plants apply more broadly: the risk of herbivore outbreaks is elevated whenever plants with low-enough herbivore loads can perfectly compensate or overcompensate for losses to herbivory.

Herbivore population dynamics in response to plant allocation strategies

When herbivores feed, plants may respond by altering the quantity of edible biomass available to future feeders through mechanisms such as compensatory regrowth of edible structures or allocation of biomass to inedible reserves. Previous work showed that some forms of compensatory regrowth can drive insect outbreaks, but this work assumed regrowth occurred without any energetic cost to the plant. While this is a useful simplifying assumption for gaining preliminary insights, plants face an inherent trade-off between allocating energy to regrowth versus storage. Therefore, we cannot truly understand the role of compensatory regrowth in driving insect outbreaks without continuing on to more realistic scenarios. In this paper, we model the interaction between insect herbivores and plants that have a trade-off between compensatory regrowth and allocation to inedible reserves in response to herbivory. We found that the plant’s allocation strategy, described in our model by parameters representing the strength of the overcompensatory response and the rates at which energy is stored and mobilized for growth, strongly affects whether herbivore outbreaks occur. Additional factors, such as the strength of food limitation and herbivore interference while feeding, influence the frequency of the outbreaks. Overall, we found a possible new role of overcompensation to promote herbivore fluctuations when it co-occurs with allocation to inedible reserves. We highlight the importance of considering trade-offs between tolerance mechanisms that plants use in response to herbivory by showing that new dynamics arise when different plant allocation strategies occur simultaneously.

Growth and flowering responses of eelgrass to simulated grazing and fecal addition by Brant Geese

AbstractSeagrasses provide a wide range of ecosystem services and are prioritized in conservation planning. Management of this dynamic community requires describing seagrass responses to repeated grazing by dependent herbivores. Using two experimental randomized block designs in Humboldt Bay, California, we tested responses of eelgrass (Zostera marina) to separate and combined effects of simulated Brant Goose (Branta bernicla) grazing (clipping to a uniform height) and fecal pellet addition, and then to medium (MED) and severe (SEV) intensities of simulated Brant flock visitation. In Experiment 1, clipping with fecal addition (FC) stimulated more clonal addition of shoots than controls or clipping and fecal addition alone (standardized β = 0.26 relative to control). The FC treatment also had the largest positive, delayed effect (β = 0.135) on density of flowering shoots. In Experiment 2, the MED intensity treatment had positive effects on shoot density (β = 0.149), rates of leaf extension (β = 0.16), and leaf productivity (β = 0.20). These MED treatment responses resulted in the highest measures of above‐ground (β = 0.099) and below‐ground (β = 0.32) biomass. Whole shoot (βMED = 0.33, βSEV = −0.36) and landscape (βMED = 0.34, βSEV = −0.23) levels of productivity for the MED treatment reflected the parabolic shape predicted by the compensatory regrowth hypothesis, likely because light and nutrient resources were available to the eelgrass and meristems that were not damaged. Meristem injury in the SEV treatment likely explained why productivity was low, but the high SEV resource levels allowed these plants to recover to control levels in approximately eight to 10 weeks after the last treatment applications. The MED level of shoot and landscape productivity would optimize low intertidal accessibility for Brant to the most nutritiously valuable leaves. Brant grazing and detrital pathways of carbon flow are likely linked with Brant‐induced productivity contributing to the detrital pathway when Brant are present and after the birds leave the estuary. The effect of Brant grazing on eelgrass sexual reproduction may increase long‐term resilience of estuarine ecosystems.

Oxygenation pattern and compensatory responses to hypoxia and hypercapnia following bilateral carotid body resection in humans.

Five years after bilateral carotid body resection (bCBR) performed in four patients, the absence of the hypoxic ventilatory response persisted, suggesting no compensatory regrowth. Breathing hypoxic gas mixtures (15% and 12%) results in a lower (by ∼10%) minimal blood oxygen saturation ( ) in bCBR patients compared to heart failure subjects (CHF) with intact peripheral chemoreceptors. After bCBR, patients were characterized by a greater short-term variability in during mild hypoxia in comparison to the CHF group. The ventilatory response to hypercapnia was preserved following bCBR and was sufficient to maintain minimal at levels comparable to controls when combined with hypoxia. Bilateral CBR - a novel treatment modality for sympathetically mediated diseases - should be used with caution due to the risk of significant desaturation even during mild hypoxia equivalent to that experienced during long-haul air travel and high altitude. Carotid body resection has been proposed as a novel treatment for sympathetically mediated diseases but the safety of bilateral carotid body resection (bCBR) for blood oxygenation during hypoxic stress (long-haul flights or high altitude) remains uncertain. Also unknown is whether central ventilatory drive is sufficient to maintain adequate oxygen saturation when exposed to hypercapnia with concomitant hypoxia. Thus, we administered: 15% O2 , 12% O2 , 5% CO2 /12% O2 and 5% CO2 /95% O2 to a group of four patients with congestive heart failure (65±2.9years) in whom bCBR was performed 5years earlier. Ventilatory, haemodynamic and blood oxygen saturation ( ) responses were recorded non-invasively and compared to control groups with intact peripheral chemoreceptors (both healthy and heart failure patients). First, we confirmed that the ventilatory response to hypoxia was eliminated in patients with bCBR, although the increase in cardiac output was preserved. Second, administration of hypoxic gas mixtures resulted in a larger decrease in and greater short-term variability of the leading to a lower minimal for both hypoxia levels in the bCBR group compared to heart failure controls (82.5±1.2% vs. 91.6±2.3% for 15% O2 and 73.8±4.0% vs. 83.7±3.1% for 12% O2 ). Third, in bCBR patients the ventilatory response to hypercapnia was present and sufficient to maintain a minimal at a level comparable to heart failure controls following administration of 5% CO2 /12% O2 (88.7±4.2% vs. 91.1±2.8%). We conclude that bCBR carries a risk of significant oxygen desaturation even during mild hypoxia. Despite preservation of central chemosensitivity, future studies should focus on unilateral CBR or on pharmacological modulation of peripheral chemosensitivity.

Myofibroblast contraction is essential for generating and regenerating the gas-exchange surface.

A majority (~95%) of the gas-exchange surface area is generated through septa formation during alveologenesis. Disruption of this process leads to alveolar simplification and bronchopulmonary dysplasia (BPD), a prevalent disorder in premature infants. Although several models have been proposed, the mechanism of septa formation remains under debate. Here we show that inactivation of myosin light chain kinase (MLCK), a key factor required for myofibroblast contraction, disrupted septa formation, supporting the myofibroblast contraction model of alveologenesis. The alveoli simplification phenotype was accompanied by decreased yes-associated protein (YAP), a key effector in the Hippo mechanotransduction pathway. Expression of activated YAP in Mlck-mutant lungs led to partial reversal of alveolar simplification. In the adult, although Mlck inactivation did not lead to simplification, it prevented reseptation during compensatory regrowth in the pneumonectomy model. These findings revealed that myofibroblast reactivation and contraction are requisite steps toward regenerating the gas-exchange surface in diseases such as BPD and chronic obstructive pulmonary disease (COPD).

Compensatory regrowth of the mouse bladder after partial cystectomy.

Cystectomy is the removal of all or part of the urinary bladder. It has been observed that there is significant regrowth of the bladder after partial cystectomy and this has been proposed to be through regeneration of the organ. Regrowth of tissue in mammals has been proposed to involve compensatory mechanisms that share many characteristics of true regeneration, like the growth of specialized structures such as blood vessels or nerves. However, the overall structure of the normal organ is not achieved. Here we tested if bladder growth after subtotal cystectomy (STC, removal of 50% of the bladder) was compensatory or regenerative. To do this we subjected adult female mouse bladders to STC and assessed regrowth using several established cellular parameters including histological, gene expression, cytokine accumulation and cell proliferation studies. Bladder function was analyzed using cystometry and the voiding stain on paper (VSOP) technique. We found that STC bladders were able to increase their ability to hold urine with the majority of volume restoration occurring within the first two weeks. Regenerating bladders had thinner walls with less mean muscle thickness, and they showed increased collagen deposition at the incision as well as throughout the bladder wall suggesting that fibrosis was occurring. Cell populations differed in their response to injury with urothelial regeneration complete by day 7, but stromal and detrusor muscle still incomplete after 8wks. Cells incorporated EdU when administered at the time of surgery and tracing of EdU positive cells over time indicated that many newborn cells originate at the incision and move mediolaterally. Basal urothelial cells and bladder mesenchymal stem cells but not smooth muscle cells significantly incorporated EdU after STC. Since anti-inflammatory cytokines play a role in regeneration, we analyzed expressed cytokines and found that no anti-inflammatory cytokines were present in the bladder 1wk after STC. Our findings suggest that bladder regrowth after cystectomy is compensatory and functions to increase the volume that the bladder can hold. This finding sets the stage for understanding how the bladder responds to cystectomy and how this can be improved in patients after suffering bladder injury.

Fuelwood extraction intensity drives compensatory regrowth in African savanna communal lands

AbstractWoody biomass remains the primary energy source for domestic use in the developing world, raising concerns about woodland sustainability. Yet woodland regenerative capacity and the adaptive response of harvesters to localised fuelwood shortages are often underestimated or unaccounted for in fuelwood supply–demand models. Here, we explore the rates and patterns of height‐specific woody vegetation structural dynamics in three communal lands in a semiarid savanna in South Africa. Using repeat, airborne light detection and ranging, we measured height‐specific change in woody vegetation structure, and the relative influence of geology, fire, and ease of access to fuelwood. Monitoring 634,284 trees canopies over 4 years revealed high compensatory growth, particularly in the high wood extraction communal land: 34.1% of trees increased in height >1 m. Vegetation structural patterns were associated with ease of access to the communal land but were mediated by wood extraction intensity. In these communal lands, vegetation structural dynamics show rapid woody thickening as a response to repeat harvesting. However, loss of height in vegetation structure did not follow a gradient of wood extraction intensity. We propose a conceptual framework to better understand change in vegetation structural metrics and the paradoxical phenomenon of high growth in high wood extraction scenarios. We also show coadaptive responses of humans and woody vegetation to fuelwood harvesting in human–environment systems through patterns of regrowth response relative to ease of access to fuelwood resources.

Functional traits partially mediate the effects of chronic anthropogenic disturbance on the growth of a tropical tree.

Understanding how trees mediate the effects of chronic anthropogenic disturbance is fundamental to developing forest sustainable management strategies. The role that intraspecific functional diversity plays in such process is poorly understood. Several tree species are repeatedly defoliated at large scale by cattle breeders in Africa to feed livestock. In addition, these tree species are also debarked for medicinal purposes. These human-induced disturbances lead to biomass loss and subsequent decline in the tree growth. The main objective of this work is to investigate how functional traits mediate tree response to chronic anthropogenic disturbance. We used a unique data set of functional traits and growth rate of 503 individual tree of Afzelia africana. We collected data on leaf mass per area (LMA), wood density (WD) and growth rate, and recorded history of human disturbances (debarking, pruning) on individual tree from 12 populations of A. africana distributed in two ecological zones in Benin (West Africa). We tested the effect of disturbances on absolute growth rate across ontogenetic stages, assessed the role of intraspecific trait variability on growth and tested the role of tree functional strategy on the tree growth response to debarking and pruning. We found that debarking did not affect stem growth, suggesting a fast compensatory regrowth of bark wounded. Moreover, tree response to debarking was independent of the functional strategy. By contrast, we found that pruning reduced tree absolute growth; however, trees with low WD were more strongly affected by pruning than trees with high WD. Our results emphasize the importance for plant functioning of the interplay between the availability of leaves for resource acquisition and a resilience strategy by mobilizing stored resources in stem wood to be reinvested for growth under severe disturbances.

Refoliation of deciduous canopy trees following severe insect defoliation: comparison of Fagus crenata and Quercus crispula

Deciduous trees can survive severe defoliation by herbivores and often refoliate in the same season. Refoliation following severe defoliation represents compensatory regrowth to recover foliage biomass. Although the relationship between defoliation intensity and degree of refoliation at the individual level has been quantified following artificial defoliation for saplings and small trees, no study has examined the relationship for canopy trees and interspecific differences in this relationship. In this study, defoliation by gypsy moths in an outbreak year and subsequent refoliation were visually surveyed for canopy trees of Fagus crenata (n = 80) and Quercus crispula (n = 113) in central Japan. Defoliation and refoliation estimates were scored in 10% classes as the ratio to foliage present before defoliation. The degree of refoliation and the proportion of refoliated trees were high in severely defoliated trees. For 60 and 100% defoliated trees, respective refoliations were 2 and 66% for F. crenata, and 37 and 88% for Q. crispula. All of the 90 and 100% defoliated trees refoliated. These results indicate that severely defoliated trees show an increased need for refoliation to maintain metabolism. Beta regression analysis showed that Q. crispula possessed higher refoliation capability than F. crenata. This is likely associated with the relatively large storage reserves and recurrent growth flush pattern of oak species, which are strong characteristics of oaks and adaptive for response to herbivory and catastrophic disturbances. Interspecific differences in refoliation capability may exert differential effects on forest ecosystem processes, such as influencing the growth of understory species.

Wisp2 disruption represses Cxcr4 expression and inhibits BMSCs homing to injured liver.

Liver regeneration/repair is a compensatory regrowth following acute liver failure, and bone marrow-derived mesenchyme stem cell (BMSC) transplantation is an effective therapy that promotes liver regeneration/repair. Wnt1 inducible signaling pathway protein 2 (Wisp2) is highly expressed in BMSCs, however, its function remains unclear. In this work, we used clustered regularly interspaced short palindromic repeats (CRISPR)-associated protein -9 nuclease (CRISPR/Cas9) genome editing technology to knockdown Wisp2 in BMSCs, and these modified cells were then transplanted into rats which were induced by the 2-AAF/PH. By linking the expression of Cas9 to green fluorescent protein (GFP), we tracked BMSCs in the rats. Disruption of Wisp2 inhibited the homing of BMSCs to injured liver and aggravated liver damage as indicated by remarkably high levels of ALT and AST. Moreover, the key factor in BMSC transplantation, C-X-C chemokine receptor type 4 (Cxcr4), was down-regulated in the Wisp2 depleted BMSCs and had a lower expression in the livers of the corresponding rats. By tracing the GFP marker, more BMSCs were observed to differentiate into CD31 positive endothelial cells in the functional Wisp2 cells but less in the Wisp2 gene disrupted cells. In summary, Wisp2 promotes the homing of BMSCs through Cxcr4 related signaling during liver repair in rats.

Evidence that a herbivore tolerance response affects selection on floral traits and inflorescence architecture in purple loosestrife (Lythrum salicaria).

The study of the evolution of floral traits has generally focused on pollination as the primary driver of selection. However, herbivores can also impose selection on floral traits through a variety of mechanisms, including florivory and parasitism. Less well understood is whether floral and inflorescence architecture traits that influence a plant's tolerance to herbivory, such as compensatory regrowth, alter pollinator-mediated selection. Because herbivore damage to Lythrum salicaria meristems typically leads to an increase in the number of inflorescences and the size of the floral display, an experiment was conducted to test whether simulated herbivory (i.e. clipping the developing meristem) could alter the magnitude or direction of pollinator-mediated selection on a suite of floral and inflorescence architecture traits. Using a pollen supplementation protocol, pollen limitation was compared in the presence and absence of meristem damage in order to quantify any interaction between pollinator and herbivore-mediated selection on floral traits. Surprisingly, in spite of an obvious impact on floral display and architecture, with clipped plants producing more inflorescences and more flowers, there was no difference in pollen limitation between clipped and unclipped plants. Correspondingly, there was no evidence that imposing herbivore damage altered pollinator-mediated selection in this system. Rather, the herbivory treatment alone was found to alter direct selection on floral display, with clipped plants experiencing greater selection for earlier flowering and weaker selection for number of inflorescences when compared with unclipped plants. These findings imply that herbivory on its own can drive selection on plant floral traits and inflorescence architecture in this species, even more so than pollinators. Specifically, herbivory can impose selection on floral traits if such traits influence a plant's tolerance to herbivory, such as through the timing of flowering and/or the compensatory regrowth response.

Rhizophora stylosa prop roots even when damaged prevent wood-boring teredinids from toppling the trees

This study examines the ability of live Rhizophora stylosa prop roots to heal damaged tissues and defend against herbivorous attack from teredinids in three mangrove forests. Sites were chosen because damaged roots and teredinid activity were frequent. Responses of 81 R. stylosa roots to three levels of experimental damage were investigated: superficial, moderate and severe. Tannin intensity post damage was analysed using ImageJ. Losses of tissue and numbers of teredinid tunnels within damaged roots among sites were not significant in magnitude. Tissue regrowth varied significantly among roots; moderately damaged roots had an over compensatory regrowth of tissue. Yet, few roots with severe damage demonstrated the same level of excessive tissue regrowth, and many roots lost tissue to necrosis and teredinid attack. Superficially damaged roots did not succumb to teredinid attack. Rhizophora stylosa roots are able to defend against teredinid larval settlement by production of tannins in damaged cortex tissue and by an over compensatory regrowth. This study highlights the resilience and ability of mangroves to heal damaged roots and defend against woodborers. But, when losses of tissues expose the vascular cylinder, teredinid larvae will settle and tunnel into the root. The roots are then open to infection, and cell necrosis.

The Effects of Plant Compensatory Regrowth and Induced Resistance on Herbivore Population Dynamics.

Outbreaks of herbivorous insects are detrimental to natural and agricultural systems, but the mechanisms driving outbreaks are not well understood. Plant responses to herbivory have the potential to produce outbreaks, but long-term effects of plant responses on herbivore dynamics are understudied. To quantify these effects, we analyze mathematical models of univoltine herbivores consuming annual plants with two responses: (1) compensatory regrowth, which affects herbivore survival in food-limited situations by increasing the amount of food available to the herbivore; and (2) induced resistance, which reduces herbivore survival proportional to the strength of the response. Compensatory regrowth includes tolerance, where plants replace some or all of the consumed biomass, and overcompensation, where plants produce more biomass than was consumed. We found that overcompensation can cause bounded fluctuations in the herbivore density (called outbreaks here) by itself, whereas neither tolerance nor induced resistance can cause an outbreak on its own. Food limitation and induced resistance can also drive outbreaks when they act simultaneously. Tolerance damps these outbreaks, but overcompensation, by contrast, qualitatively changes the conditions under which the outbreaks occur. Not properly accounting for these interactions may explain why it has been difficult to document plant-driven insect outbreaks and could undermine efforts to control herbivore populations in agricultural systems.

Variation in GmAOS1 promoter is associated with soybean defense against insect attack

Soybean (Glycine max L.) displays considerable variation in its resistance to insects. Characterization and application of insect resistance genes in soybean germplasm may be a sustainable and economical approach to soybean production. In this study, we demonstrated that the gene encoding allene oxide synthase (GmAOS1) was associated with soybean defense against the common cutworm (CCW, Spodoptera litura Fabricius) attack through gene expression analysis and association analysis. The expression of GmAOS1 in stems and leaves of resistant accessions was higher than that of susceptible accessions in the undamaged soybean plant. After CCW feeding, the transcript levels of GmAOS1 increased and reached their maximal values sooner in the leaves of resistant accession than they did in susceptible accession. The genomic sequence of GmAOS1 was re-sequenced and aligned in 184 soybean accessions. Forty-three SNPs/Indels were found in its promoter region and only one single nucleotide polymorphism was found in its coding region. Association analysis showed that GmAOS1 promoter polymorphisms were significantly associated with three soybean defense indexes. GmAOS1_19 was the most resistant allele associated with soybean resistance to CCW. Two elements (TATCCAT/C motif and Box II-like sequence) located at genomic regions upstream of GmAOS1_21 were significantly associated with compensatory regrowth of soybean plants. Gene expression analysis indicated that the accessions containing different GmAOS1 promoters and with different CCW resistance grades showed different expression levels of GmAOS1. Our data indicate that variation in GmAOS1 promoter is associated with soybean defense against CCW attack.

Loss of specificity: native but not invasive populations of Triadica sebifera vary in tolerance to different herbivores

During introduction, invasive plants can be released from specialist herbivores, but may retain generalist herbivores and encounter novel enemies. For fast-growing invasive plants, tolerance of herbivory via compensatory regrowth may be an important defense against generalist herbivory, but it is unclear whether tolerance responses are specifically induced by different herbivores and whether specificity differs among native and invasive plant populations. We conducted a greenhouse experiment to examine the variation among native and invasive populations of Chinese tallow tree, Triadica sebifera, in their specificity of tolerance responses to herbivores by exposing plants to herbivory from either one of two generalist caterpillars occurring in the introduced range of Triadica. Simultaneously, we measured the specificity of another defensive trait, extrafloral nectar (EFN) production, to detect potential tradeoffs between resistance and tolerance of herbivores. Invasive populations had higher aboveground biomass tolerance than native populations, and responded non-specifically to either herbivore, while native populations had significantly different and specific aboveground biomass responses to the two herbivores. Both caterpillar species similarly induced EFN in native and invasive populations. Plant tolerance and EFN were positively correlated or had no relationship and biomass in control and herbivore-damaged plants was positively correlated, suggesting little costs of tolerance. Relationships among these vegetative traits depended on herbivore type, suggesting that some defense traits may have positive associations with growth-related processes that are differently induced by herbivores. Importantly, loss of specificity in invasive populations indicates subtle evolutionary changes in defenses in invasive plants that may relate to and enhance their invasive success.

Effects of insect herbivory on induced chemical defences and compensation during early plant development in Penstemon virgatus.

The lack of studies assessing the simultaneous expression of tolerance and resistance traits during seedling development and overall seedling defences as compared with adult plants, in general, constitutes a significant research need that can greatly improve our understanding of overall investment in defences during plant ontogeny. Using two seedling and two juvenile stages of the perennial herb Penstemon virgatus (Plantaginaceae) evaluations were made of (a) patterns of investment in constitutive chemical defences [i.e. iridoid glycosides (IGs)], and (b) simultaneous variation in the short-term ability of seedling and juvenile stages to induce resistance traits, measured as induced chemical defences, or tolerance traits, measured as compensatory re-growth following moderate levels of damage by a specialist insect herbivore. Plants were highly defended during most of their transition from seedling to early juvenile stages, reaching a constant approx. 20 % dry weight total IGs. Furthermore, following 30 % above-ground tissue damage, seedlings and juvenile stages were equally able to induce resistance, by raising their IG concentration by approx. 8 %, whereas compensatory re-growth was only achieved at young juvenile but not seedling stages. Two major trends emerged from this study: (1) in contrast to expected and previously observed trends, in this perennial plant species, seedlings seem to be one of the most well-defended stages as compared with adult ones; (2) high levels of constitutive defences did not limit the ability of young developmental stages to induce resistance following damage, although this response may come with a cost (i.e. decreased compensation) in young seedling stages. Hence, as has been previously demonstrated in few other systems, these results points towards an indirect evidence for a trade-off between tolerance and resistance traits at some, but not all, developmental stages; making them often difficult to detect.

Interspecific Variation in Compensatory Regrowth to Herbivory Associated with Soil Nutrients in Three Ficus (Moraceae) Saplings

Plant compensatory regrowth is an induced process that enhances plant tolerance to herbivory. Plant behavior against herbivores differs between species and depends on resource availability, thus making general predictions related to plant compensatory regrowth difficult. To understand how soil nutrients determine the degree of compensatory regrowth for different plant species, we selected saplings of three Ficus species and treated with herbivore insects and artificial injury in both glasshouse conditions and in the field at two soil nutrient levels. Compensatory regrowth was calculated by biomass, relative growth rate and photosynthetic characteristics. A similar pattern was found in both the glasshouse and in the field for species F. hispida, where overcompensatory regrowth was triggered only under fertile conditions, and full compensatory regrowth occurred under infertile conditions. For F. auriculata, overcompensatory regrowth was stimulated only under infertile conditions and full compensatory regrowth occurred under fertile conditions. Ficus racemosa displayed full compensatory regrowth in both soil nutrient levels, but without overcompensatory regrowth following any of the treatments. The three Ficus species differed in biomass allocation following herbivore damage and artificial injury. The root/shoot ratio of F. hispida decreased largely following herbivore damage and artificial injury, while the root/shoot ratio for F. auriculata increased against damage treatments. The increase of shoot and root size for F. hispida and F. auriculata, respectively, appeared to be caused by a significant increase in photosynthesis. The results indicated that shifts in biomass allocation and increased photosynthesis are two of the mechanisms underlying compensatory regrowth. Contrasting patterns among the three Ficus species suggest that further theoretical and empirical work is necessary to better understand the complexity of the plant responses to herbivore damage.

Plant Induced Defenses Depend More on Plant Age than Previous History of Damage: Implications for Plant-Herbivore Interactions

Herbivore-induced plant responses can significantly change as a function of plant developmental stage and previous history of damage. Yet, empirical tests that assess the combined role of multiple damage events and age-dependent constraints on the ability of plants to induce defenses within and among tissues are scarce. This question is of particular interest for annual and/or short-lived perennial plant species, whose responses to single or multiple damage events over a growing season are likely to interact with ontogenetic constraints in affecting a plant's ability to respond to herbivory. Using Plantago lanceolata and one of its specialist herbivores, Junonia coenia, we examined the effect of plant ontogeny (juvenile vs. mature developmental stages) and history of damage (single and multiple damage events early and/or late in the season) on plant responses to leaf damage. Plant responses to herbivory were assessed as induced chemical defenses (iridoid glycosides) and compensatory regrowth, in both above- and below-ground tissues. We found that constitutive concentration of iridoid glycosides markedly increased as plants matured, but plant ability to induce chemical defenses was limited to juvenile, but not mature, plant stages. In addition, induced defenses observed 7d following herbivory in juvenile plants disappeared 5wk after the first herbivory event, and mature plants that varied considerably in the frequency and intensity of damage received over 5wk, did not differ significantly in their levels of chemical defenses. Also, only small changes in compensatory regrowth were detected. Finally, we did not observe changes in below-ground tissues' defenses or biomass a week following 50% removal of leaf tissues at either age class or history of damage. Together, these results suggest that in P. lanceolata and perhaps other systems, ontogenetic trajectories in plant growth and defenses leading to strong age-dependent induced responses may prevail over herbivore-induced indirect interactions.

Multiple common garden experiments suggest lack of local adaptation in an invasive ornamental plant

Aims Adaptive evolution along geographic gradients of climatic conditions is suggested to facilitate the spread of invasive plant species, leading to clinal variation among populations in the introduced range. We investigated whether adaptation to climate is also involved in the invasive spread of an ornamental shrub, Buddleja davidii, across western and central Europe. Methods We combined a common garden experiment, replicated in three climatically different central European regions, with reciprocal transplantation to quantify genetic differentiation in growth and reproductive traits of 20 invasive B. davidii populations. Additionally, we compared compensatory regrowth among populations after clipping of stems to simulate mechanical damage.

Delayed flowering as a potential benefit-decreasing cost of compensatory regrowth

Some monocarpic plants show surprising variation in response to natural and simulated grazing. In favourable conditions, injured plants may grow larger and produce more fruits and viable seeds compared with their uninjured counterparts. Such overcompensation, however, is associated with potential costs, particularly delays in flowering and fruit maturation. These delays may further increase a risk of frost injury before seed maturation in the early autumn. We tested the effects of artificially advanced night frosts on the compensatory capacity (i.e., the capacity to compensate the lost biomass by regrowth) of the monocarpic herb Erysimum strictum P. Gaertn., B. Mey., and Scherb. in a common garden experiment. Two simulated herbivory treatments were applied: 25% apical clipping in early vs. late June. Frost treatment was applied in a climate chamber in late August – early September, about 3 weeks before the first natural frost period. Apically damaged plants not exposed to frost treatment produced 1.9–2.6 times more total biomass and 2.5 times more fruit than intact plants. Frost treatment reduced plant performance by 35%–48%, but in contrast to our expectation, there was no significant difference between intact and apically damaged plants in response to early frost. In spite of the delay in phenology, compensatory regrowth did not increase the risk of frost injury. We conclude that while early night frosts imply a potential risk to monocarpic herbs recovering from herbivory, possibly other suboptimal conditions, such as drought in late summer, may provide a greater threat for early-flowering meadow and grassland plants recovering from grazing. Possibly multiple selective forces and environmental risk factors operate together in the evolution of grazing tolerance associated with flowering phenology, plant growth strategy, and architecture.