Volatile communication between plants that affects herbivory: a meta‐analysis

Volatile communication between plants causing enhanced defence has been controversial. Early studies were not replicated, and influential reviews questioned the validity of the phenomenon. We collected 48 well‐replicated studies and found overall support for the hypothesis that resistance increased for individuals with damaged neighbours. Laboratory or greenhouse studies and those conducted on agricultural crops showed stronger induced resistance than field studies on undomesticated species, presumably because other variation had been reduced. A cumulative analysis revealed that early, non‐replicated studies were more variable and showed less evidence for communication. Effects of habitat and plant growth form were undetectable. In most cases, the mechanisms of resistance and alternative hypotheses were not considered. There was no indication that some response variables were more likely to produce large effects. These results indicate that plants of diverse taxonomic affinities and ecological conditions become more resistant to herbivores when exposed to volatiles from damaged neighbours.     

Long-term grazing affects relationships between nitrogen form uptake and biomass of alpine meadow plants

Trade-off between nutrient uptake rate and product accumulation has been found among species characterized as acquisitive and conservative strategies in resource utilization. However, long-term grazing causes changes in soil nutrient availability and plant species abundance by selective foraging and resource allocation between above- and belowground organs, which may cover up such trade-off. However, little is known whether the trade-off can be observed among species in community without grazing disturbance, and how grazing influences the trade-off. We conducted a ¹⁵N labelling experiment in winter grazing and grazing release alpine meadow communities on the Tibet Plateau. We examined changes in N form uptake of 11 common species and relationship of N chemical uptake rate with aboveground biomass. Grazing release increased soil \({\text{NH}}_{4}^{ + }\)–N and \({\text{NO}}_{3}^{ - }\)–N, and increased \({\text{NO}}_{3}^{ - }\)–N uptake rate in two species and \({\text{NH}}_{4}^{ + }\)–N uptake rate in three species. Meanwhile, grazing release decreased aboveground biomass of three species and two of them belong to those species’ increased N uptake rate. Contrarily, grazing release increased aboveground biomass of four species and none belongs to the changed N uptake rate. Thus, grazing release caused changes of plant nutrient uptake rate and aboveground production in different directions, which explains the negative relationship of N uptake rate with aboveground biomass in ungrazed community. Our results indicate that the increase in nutrient uptake is probably one of the mechanisms for acquisitive species to cope with the raising nutrient availability and/or competition from the conservative dominant grasses after grazing release.     

How geitonogamous selfing affects sex allocation in hermaphrodite plants

Does the mode of self-pollination affect the evolutionarily stable allocation to male vs. female function? We distinguish the following scenarios. (1) An ‘autogamous’ species, in which selfing occurs within the flower prior to opening. The pollen used in selfing is a constant fraction of all pollen grains produced. (2) A species with ‘abiotic pollination’, in which selfing occurs when pollen dispersed in one flower lands on the stigma of a nearby flower on the same plant (geitonogamy). The selfing rate increases with male allocation but a higher selfing rate does not mean a reduced export of pollen. (3) An ‘animal-pollinated’ species with geitonogamous selfing. Here the selfing rate also increases with male allocation, but pollen export to other plants in the population is a decelerating function of the number of simultaneously open flowers. In all three models selfing selects for increased female allocation. For model 3 this contradicts the general opinion that geitonogamous selfing does not affect evolutionarily stable allocations. In all models, the parent benefits more from a female-biased allocation than any other individual in the population. In addition, in models 2 and 3, greater male allocation results in more local mate competition. In model 3 and in model 2 with low levels of inbreeding depression, hermaphroditism is evolutionarily stable. In model 2 with high inbreeding depression, the population converges to a fitness minimum for the relative allocation to male function. In this case the fitness set is bowed inwards, corresponding with accelerating fitness gain curves. If the selfing rate increases with plant size, this is a sufficient condition for size-dependent sex allocation (more allocation towards seeds in large plants) to evolve. We discuss our results in relation to size-dependent sex allocation in plants and in relation to the evolution of dioecy.     

Resource capture, biomass allocation and growth in herbaceous plants

Plant species differ widely in their rate of biomass production, even when grown under optimal conditions. A key question concerns the extent to which these growth rates correlate with the uptake of carbon and nitrogen and with the biomass allocation between leaves and roots. Recent data show that the answer to this question differs for mono- and dicotyledons, and that more than biomass allocation, it is the ratio between the activities of leaves and roots that correlates with the growth rate of a plant.     

Volatile organic compounds released by maize following herbivory or insect extract application and communication between plants

To protect themselves from herbivory, plants have evolved an arsenal of physical and chemical defences and release a variety of volatile organic compounds (VOCs). By releasing these VOCs, a signalling plant can both reduce herbivory, sometimes by more than 90%, and also warn neighbouring plants about an attack. The aim of this study was to assess the influence of herbivory and insect extract application on VOC release by damaged/treated and nearby undamaged/untreated maize plants. We confirmed that European corn borer (Ostrinia nubilalis) larvae attack or larvae extract application induced maize VOC release. Greater amounts of (Z)‐3‐hexenal, (E)‐2‐hexenal, (Z)‐3‐hexen‐1‐ol, (E)‐2‐hexen‐1‐ol, β‐myrcene, (Z)‐3‐hexen‐1‐yl acetate, 1‐hexyl acetate, (Z)‐ocimene, linalool, benzyl acetate, methyl salicylate, indole, methyl anthranilate, geranyl acetate, β‐caryophyllene, (E)‐β‐farnesene and (Z)‐3‐hexenal, (Z)‐3‐hexen‐1‐ol, (Z)‐3‐hexen‐1‐yl acetate, (Z)‐ocimene, linalool, indole, methyl anthranilate, geranyl acetate, β‐caryophyllene and (E)‐β‐farnesene were released as a result of biotic stress after insect attack or insect extract application. The amounts of each VOC released were qualitatively and quantitatively distinct and dependent on time after biotic stress exposure. However, for all biotic stresses, significantly lower VOC induction was measured when leaves were damaged/treated for three days, as compared to seven days. Our work also demonstrated that undamaged/untreated neighbouring plants also release significant amounts of VOCs. This suggests that VOC emission by a damaged/treated plant stimulates VOC induction in nearby undamaged/untreated plants. However, the concentrations of all VOCs released by neighbouring undamaged/untreated maize plants were lower than those from damaged/treated plants and were negatively correlated with distance from a damaged/treated plant. Still, significant VOC induction occurred in undamaged/untreated plants even at 3 m distance from a damaged/infected plant. Our work suggests that maize plant protective defence responses (VOC emission) can be induced via application of European corn borer extracts.     

Volatile communication among sagebrush branches affects herbivory: timing of active cues

Airborne communication can affect systemic induced resistance to herbivory on neighboring branches and individual plants. Sagebrush is the best known example of this phenomenon although the mechanisms of this communication system remain unidentified. We do not know the timing of emission or the chemical nature of the active cue. We investigated the timing of this phenomenon by using plastic bags to prevent propagation of volatile compounds and experimentally manipulated the timing of removal of these bags. We found that blocking the volatiles prevented systemic induced resistance. Experimentally allowing clipped branches to release cues for up to 3 days after clipping caused a reduction in damage in neighboring branches on the clipped plants. This indicates that active cues are released from the time we clipped for the next 3 days or that cues released immediately remained active over this time period. As we continue to evaluate potential chemicals as active cues in plant communication, this prolonged effectiveness may provide an important screen against which to evaluate any putative signals. Handling editor: Robert Glinwood     

Sexual differences in biomass and nutrient allocation of first-year Silene dioica plants

Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences. Received: 07 May 1998 / Accepted: 30 October 1998     

Visibility vs. biomass in flowers: exploring corolla allocation in Mediterranean entomophilous plants

Background and Aims

While pollinators may in general select for large, morphologically uniform floral phenotypes, drought stress has been proposed as a destabilizing force that may favour small flowers and/or promote floral variation within species.

Methods

The general validity of this concept was checked by surveying a taxonomically diverse array of 38 insect-pollinated Mediterranean species. The interplay between fresh biomass investment, linear size and percentage corolla allocation was studied. Allometric relationships between traits were investigated by reduced major-axis regression, and qualitative correlates of floral variation explored using general linear-model MANOVA.

Key Results

Across species, flowers were perfectly isometrical with regard to corolla allocation (i.e. larger flowers were just scaled-up versions of smaller ones and vice versa). In contrast, linear size and biomass varied allometrically (i.e. there were shape variations, in addition to variations in size). Most floral variables correlated positively and significantly across species, except corolla allocation, which was largely determined by family membership and floral symmetry. On average, species with bilateral flowers allocated more to the corolla than those with radial flowers. Plant life-form was immaterial to all of the studied traits. Flower linear size variation was in general low among conspecifics (coefficients of variation around 10 %), whereas biomass was in general less uniform (e.g. 200–400 mg in Cistus salvifolius). Significant among-population differences were detected for all major quantitative floral traits.

Conclusions

Flower miniaturization can allow an improved use of reproductive resources under prevailingly stressful conditions. The hypothesis that flower size reflects a compromise between pollinator attraction, water requirements and allometric constraints among floral parts is discussed.Key words: Allometry, biomass, corolla, drought, evolution, flower, Mediterranean, sclerophyllous, size, variation, water     

Red:far-red light conditions affect the emission of volatile organic compounds from barley (Hordeum vulgare), leading to altered biomass allocation in neighbouring plants

Background and Aims Volatile organic compounds (VOCs) play various roles in plant–plant interactions, and constitutively produced VOCs might act as a cue to sense neighbouring plants. Previous studies have shown that VOCs emitted from the barley (Hordeum vulgare) cultivar ‘Alva’ cause changes in biomass allocation in plants of the cultivar ‘Kara’. Other studies have shown that shading and the low red:far-red (R:FR) conditions that prevail at high plant densities can reduce the quantity and alter the composition of the VOCs emitted by Arabidopsis thaliana, but whether this affects plant–plant signalling remains unknown. This study therefore examines the effects of far-red light enrichment on VOC emissions and plant–plant signalling between ‘Alva’ and ‘Kara’.Methods The proximity of neighbouring plants was mimicked by supplemental far-red light treatment of VOC emitter plants of barley grown in growth chambers. Volatiles emitted by ‘Alva’ under control and far-red light-enriched conditions were analysed using gas chromatography–mass spectrometry (GC-MS). ‘Kara’ plants were exposed to the VOC blend emitted by the ‘Alva’ plants that were subjected to either of the light treatments. Dry matter partitioning, leaf area, stem and total root length were determined for ‘Kara’ plants exposed to ‘Alva’ VOCs, and also for ‘Alva’ plants exposed to either control or far-red-enriched light treatments.Key Results Total VOC emissions by ‘Alva’ were reduced under low R:FR conditions compared with control light conditions, although individual volatile compounds were found to be either suppressed, induced or not affected by R:FR. The altered composition of the VOC blend emitted by ‘Alva’ plants exposed to low R:FR was found to affect carbon allocation in receiver plants of ‘Kara’.Conclusions The results indicate that changes in R:FR light conditions influence the emissions of VOCs in barley, and that these altered emissions affect VOC-mediated plant–plant interactions.     

Nitrogen addition affects plant biomass allocation but not allometric relationships among different organs across the globe

全球尺度上氮添加影响植物生物量分配但不影响不同器官间的异速生长关系 生物量在不同器官间的分配是植物对环境变化响应的一个关键生态生理学过程。然而,在全球尺度上有关不同陆地生态系统植物生物量分配对氮沉降响应的认识还比较欠缺。本文通过整合分析333篇已发表文章的5474个观测值,基于“最优分配假说”和“异速分配假说”,评估了全球尺度上氮添加对植物生物量及其在不同器官间分配的影响。结果表明：(1)氮添加显著增加了整株植物或不同器官的生物量,降低了根冠比和根质量分数,但对叶质量分数和茎质量分数无显著影响;(2)氮添加对不同器官质量分数的影响受实验条件、植物功能性状、纬度、氮添加率等因子单独或交互作用的调控;(3)氮添加对生物量在不同器官间的异速分配率无显著影响,表明氮添加导致的根冠比和根质量分数减少是在异速分配模式下由整株植物生物量增加而致。虽然氮添加改变了地上和地下部分的质量分数,但不同器官生物量间异速分配模式的稳定性表明“异速分配假说”能更好地描述植物生物量分配对氮添加的响应规律。该研究结果将有助于深入认识氮沉降环境下植物生物量的分配规律,同时表明将生物量分配纳入有关氮沉降对陆地生态系统影响的预测模型中的重要性。     

The effect of nutrient availability on biomass allocation patterns in 27 species of herbaceous plants

We investigated allocation to roots, stems and leaves of 27 species of herbaceous clonal plants grown at two nutrient levels. Allocation was analyzed as biomass ratios and also allometrically. As in other studies, the fraction of biomass in stems and, to a lesser extent, in leaves, was usually higher in the high-nutrient treatment than in the low-nutrient treatment, and the fraction of biomass in roots was usually higher under low-nutrient conditions. The relationship between the biomass of plant structures fits the general allometric equation, with an exponent 1 in most of the species. The different biomass ratios under the two nutrient conditions represented points on simple allometric trajectories, indicating that natural selection has resulted in allometric strategies rather than plastic responses to nutrient level. In other words, in most of the species that changed allocation in response to the nutrient treatment, these changes were largely a consequence of plant size. Our data suggest that some allocation patterns that have been interpreted as plastic responses to different resource availabilities may be more parsimoniously explained as allometric strategies.     

Patterns of biomass allocation to male and female functions in plants with different mating systems

Summary Using dry weight biomass we examined the patterns of investment in male and female functions (prezygotic cost) in plants with different mating systems. All the flower parts of both xenogamous and facultatively xenogamous species were heavier, i.e., larger, than those of facultatively autogamous species. Likewise, the dry weights of all the flower parts of xenogamous species exceeded those of facultatively xenogamous species. On a relative basis, xenogamous species invested less in calyces and more in corollas compared to species with the other mating systems. Facultatively autogamous species invested relatively more in pistils. Xenogamous species invested relatively more in stamens than do facultatively autogamous species. The ratios of dry weight stamens to dry weight pistils were equivalent in xenogamous and facultatively xenogamous species.The available data from xenogamous species suggest a pattern of resource allocation that is independent of sexual system (perfect-flowered, monoecious, or dioecious) and pollen vector. The cost of mating (prezygotic cost) was male biased and frequently exceeded by parental investment (postzygotic cost). These results are not consistent with models that predict equal allocation of resources to male and female sexual function but are consistent with those that predict unequal allocation of resources to those functions in outbreeding hermaphroditic angiosperms. Two additional lines of evidence are inconsistent with the expectations of sex allocation theory. First, resource allocation to sexual function was not equal in wind-pollinated species. Second, relative amounts of the resources allocated to male vis-à-vis female function did not decrease between xenogamy and facultative xenogamy i.e., with an increase in the selfing rate.     

The relation between above- and belowground biomass allocation patterns and competitive ability

Summary In a 2-year experiment, the evergreen shrubsErica tetralix andCalluna vulgaris (dominant on nutrient-poor heathland soils) and the perennial deciduous grassMolinia caerulea (dominant on nutrient-rich heathland soils) were grown in replacement series in a factorial combination of four competition types (no competition, only aboveground competition, only belowground competition, full competition) and two levels of nutrient supply (no nutrients and 10 g N+2 g P+10 g K m⁻² yr⁻¹). Both in the unfertilized and in the fertilized treatmentsMolinia allocated about twice as much biomass to its root system than didErica andCalluna. In all three species the relative amount of biomass allocated to the roots was lower at high than at low nutrient supply. The relative decrease was larger forMolinia than forErica andCalluna. In the fertilized monocultures biomass of all three species exceeded that in the unfertilized series.Molinia showed the greatest biomass increase. In the unfertilized series no effects of interspecific competition on the biomass of each species were observed in either of the competition treatments. In the fertilized mixtures where only belowground competition was possibleMolinia increased its biomass at the expense of bothErica andCalluna. When only aboveground competition was possible no effects of interspecific competition on the biomass of the competing species were observed. However, in contrast with the evergreens,Molinia responded by positioning its leaf layers relatively higher in the canopy. The effects of full competition were similar to those of only belowground competition, so in the fertilized series belowground competition determined the outcome of competition. The high competitive ability ofMolinia at high nutrient supply can be attributed to the combination of (1) a high potential productivity, (2) a high percentage biomass allocation to the roots, (3) an extensive root system exploiting a large soil volume, and (4) plasticity in the spatial arrangement of leaf layers over its tall canopy. In the species under study the allocation patterns entailed no apparent trade-off between the abilities to compete for above- and belowground resources. This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.     

密度制约决定的植物生物量分配格局

基于自然环境下红葱(Allium cepa var.proliferum)个体各器官生物量积累动态、生物量分配比例动态、生物量比率动态和形态性状对不同种群密度(36、49、64、121和225株·m-2)响应的模拟实验,分析了密度制约对其生物量分配格局的影响。结果表明:红葱地上部分、叶和鞘的生物量分配比例均随密度的增加而增加,地下部分和鳞茎的分配比例随密度的增加而下降,而根的分配比例未随密度发生显著变化。除根:叶、根:地上比在密度处理间无显著差异外,各器官间生物量比率均表现出明显的密度依赖性。随着个体的生长,根:鞘、根:叶、根:地上比逐渐减小,鳞茎:叶、鳞茎:鞘、鳞茎:地上比先减小后增加,而地上:地下比先增加后减小。比叶面积与密度呈显著正相关(P<0.01),叶面积和根长与密度呈显著负相关(P<0.05),而比根长不受密度的影响。由此可见,种内竞争水平会对植物体内的资源分配产生较大影响;植物生物量分配格局响应不同密度具有可塑性,随着密度的增加,红葱个体会增加地上营养器官的生物量分配,并以减小地下无性繁殖器官的生物量分配为代价。最优化分配理论仅在无竞争存在的情况下适用,当竞争发生时,种群密度及其制约性调节是决定...     

Heterogeneous light supply affects growth and biomass allocation of the understory fern Diplopterygium glaucum at high patch contrast

Spatial heterogeneity in resource supply is common and responses to heterogeneous resource supply have been extensively documented in clonal angiosperms but not in pteridophytes. To test the hypotheses that clonal integration can modify responses of pteridophytes to heterogeneous resource supply and the integration effect is larger at higher patch contrast, we conducted a field experiment with three homogeneous and two heterogeneous light treatments on the rhizomatous, understory fern Diplopterygium glaucum in an evergreen broad-leaved forest in East China. In homogeneous treatments, all D. glaucum ramets in 1.5 m×1.5 m units were subjected to 10, 40 and 100% natural light, respectively. In the heterogeneous treatment of low patch contrast, ramets in the central 0.5 m×0.5 m plots of the units were subjected to 40% natural light and their interconnected ramets in the surrounding area of the units to 100%; in the heterogeneous treatment of high patch contrast, ramets in the central plots were subjected to 10% natural light and those in the surrounding area to 100%. In the homogeneous treatments, biomass and number of living ramets in the central plots decreased and number of dead ramets increased with decreasing light supply. At low contrast heterogeneous light supply did not affect performance or biomass allocation of D. glaucum in the central plots, but at high contrast it increased lamina biomass and number of living ramets older than annual and modified biomass allocation to lamina and rhizome. Thus, clonal integration can affect responses of understory ferns to heterogeneous light supply and ramets in low light patches can be supported by those in high light. The results also suggest that effects of clonal integration depend on the degree of patch contrast and a significant integration effect may be found only under a relatively high patch contrast.