Sex ratio variation and spatial distribution of Siparuna grandiflora, a tropical dioecious shrub

Populations of dioecious plant species often exhibit biased sex ratios. Such biases may arise as a result of sex-based differences in life history traits, or as a result of spatial segregation of the sexes. Of these, sex-based differentiation in life history traits is likely to be the most common cause of bias. In dioecious species, selection can act upon the sexes in a somewhat independent way, leading to differentiation and evolution toward sex-specific ecological optima. I examined sex ratio variation and spatial distribution of the tropical dioecious shrub Siparuna grandiflora to determine whether populations exhibited a biased sex ratio, and if so, whether the bias could be explained in terms of non-random spatial distribution or sex-based differentiation in life history traits. Sex ratio bias was tested using contingency tables, a logistic regression approach was utilized to examine variation in life history traits, and spatial distributions were analyzed using Ripley's K, a second-order neighborhood analysis. I found that although populations of S. grandiflora have a male-biased sex ratio within and among years, there was no evidence of spatial segregation of the sexes. Rather, the sex ratio bias was shown to result primarily from sex-based differentiation in life history traits; males reproduce at a smaller size and more frequently than females. The sexes also differ in the relationship between plant size and reproductive frequency. Light availability was shown to affect reproductive activity in both sexes, though among infrequently flowering plants, females require higher light levels than males to flower. The results of this study demonstrate that ecologically significant sex-based differentiation has evolved in S. grandiflora. Received: 30 July 1997 / Accepted: 16 December 1997     

Simultaneous inbreeding modifies inbreeding depression in a plant–herbivore interaction

Because inbreeding is common in natural populations of plants and their herbivores, herbivore‐induced selection on plants, and vice versa, may be significantly modified by inbreeding and inbreeding depression. In a feeding assay with inbred and outbred lines of both the perennial herb, Vincetoxicum hirundinaria, and its specialist herbivore, Abrostola asclepiadis, we discovered that plant inbreeding increased inbreeding depression in herbivore performance in some populations. The effect of inbreeding on plant resistance varied among plant and herbivore populations. The among‐population variation is likely to be driven by variation in plant secondary compounds across populations. In addition, inbreeding depression in plant resistance was substantial when herbivores were outbred, but diminished when herbivores were inbred. These findings demonstrate that in plant–herbivore interactions expression of inbreeding depression can depend on the level of inbreeding of the interacting species. Furthermore, our results suggest that when herbivores are inbred, herbivore‐induced selection against self‐fertilisation in plants may diminish.     

Intersexual differences in phenology and damage by herbivores and pathogens in dioecious Rubus chamaemorus L.

Summary Shoot development and damage by herbivores and pathogens to male, female and non-floral ramets of the dioecious, perennial herb Rubus chamaemorus were studied in the field during three consecutive years. Leaves on male ramets were usually consumed more by herbivores and attacked more by fungi than were leaves on female ramets. Male ramets unfolded their leaves later than did female ramets. In 1983, when the level of herbivory was comparatively high, ramets that were fully developed in early June were more damaged by herbivores than were ramets that unfolded their leaves later. Non-floral ramets usually showed intermediate levels of damage compared to male and female flowering ramets.Defoliation caused a greater increase in ramet mortality among females than among males in a field experiment. It is suggested that the different reproductive roles of males and females differently constrain the evolution of defense against herbivores and pathogens in dioecious plants. This may result in intersexual differences both in palatability to herbivores and susceptibility to pathogens.     

Does sex matter? Gender-specificity and its influence on site-chronologies in the common dioecious shrub Juniperus communis

In recent years an increasing number of studies have shown shrubs to be reliable proxies of environmental conditions in regions where Trees − due to harsh climate conditions − are absent. Although many shrubs are monoecious, some are dioecious, which poses certain questions related to gender-specific growth as observed for trees in previous studies. Here, we address the questions whether dioecious shrubs, similar to trees, show growth differences between male and female plants, and − if so − whether this difference needs to be considered in terms of sample selection. We chose Juniperus communis. L., the most widely distributed woody plant, and a common and well-studied dioecious shrub species in the northern hemisphere, especially in the Boreal, Subarctic tundra and Alpine regions. Our samples were collected from four sites − three from the Ural Mountains and one site from Kirkenes in Norway. To see if there were differences in radial growth between sexes we performed four different analyses. First, we used multivariate explorative statistics to see if there were gender biased sub-populations and generally found no differences. Secondly, to compare growth over the lifetime of shrubs we computed cumulative annual increments of basal area which revealed no gender-specific growth patterns. Thirdly, to test if differences in radial growth between male and female shrubs affect the resulting site-chronology, we compared individual shrub chronologies with the site-chronology and found a significant differentiation between normalized correlations of gender-specific chronologies to the site-chronology. This significant difference was restricted to an overall comparison, but not evident at individual site-level. Lastly, we compared correlations of gender-specific chronologies and a mean site-chronology with monthly climate records to find only very few meaningful differences in their responses. In summary, we could not detect any clear gender-specific growth pattern in Juniperus communis but observed a trend towards more non-climatic signals in female junipers which may affect the resulting site-chronology.     

Warming affects different components of plant–herbivore interaction in a simplified community but not net interaction strength

Global warming impacts natural communities through effects on performance of individual species and through changes in the strength of interactions between them. While there is a body of evidence of the former, we lack experimental evidence on potential changes in interaction strengths. Knowledge about multispecies interactions is fundamental to understand the regulation of biodiversity and the impact of climate change on communities. This study investigated the effect of warming on a simplified community consisting of three species: rosy apple aphid Dysaphis plantaginea feeding on plantain, Plantago lanceolata, and a heterospecific neighbouring plant species, perennial ryegrass, Lolium perenne. The aphid does not feed on L. perenne. The experimental design consisted of monocultures and mixtures of L. perenne and P. lanceolata at three temperature levels. We did not find indication for indirect temperature effects on D. plantaginea through changes in leaf nitrogen or relative water content. However, experimental warming affected the life history traits of the aphid directly, in a non‐linear manner. Aphids performed best at moderate warming, where they grew faster and had a shorter generation time. In spite of the increased population growth of the aphids under warming, the herbivory rates were not changed and consequently the plant–herbivore interaction was not altered under warming. This suggests reduced consumption rates at higher temperature. Also plant competition affected the aphids but through an interaction with temperature. We provide proof‐of‐concept that net interactions between plants and herbivores should not change under warming despite direct effects of warming on herbivores when plant–plant interaction are considered. Our study stresses the importance of indirect non–trophic interactions as an additional layer of complexity to improve our understanding of how trophic interactions will alter under climate change.     

Survival costs and reproductive benefits of floral display in a sexually dimorphic dioecious shrub, Leucadendron xanthoconus

The evolutionary causes of sexual dimorphism in plants have not been as widely studied as in animals and the importance of sexual selection in causing dimorphism remains controversial. Sexual selection is most obvious when it favours the evolution of a trait which enhances mating success at the expense of decreased viability. We studied the relationship between floral display (number of inflorescences), pollinator attraction and plant survival in a dioecious shrub, Leucadendron xanthoconus. Pollinator attraction, measured as the number of insect pollinators, increased linearly with floral display in males. However, males with extravagant displays had a higher probability of dying. Our data suggest that male plants are undergoing selection on floral display for increased mating success counterbalanced by selection against plants with extravagant displays. Seed set in females did not increase with floral display, except at very low inflorescence numbers. Nor was female survival correlated with floral display. Because inflorescences are terminal in the species, selection for more inflorescences in males causes increased ramification, thinner terminal branches and smaller leaves. Thus vegetative dimorphism in this species appears to be caused by selection for extravagant floral display in males, but not females. Limits to dimorphism are imposed by survival costs of elaborate display.     

Reproductive allocation and the long-term costs of reproduction in Siparuna grandiflora, a dioecious neo-tropical shrub

1 Using a combination of observational and experimental approaches, both allocation of resources to reproduction (often called the direct cost of reproduction) and the subsequent long-term costs (the indirect, delayed or demographic cost) associated with reproductive allocation to male and female function in Siparuna grandiflora (Siparunaceae), a tropical dioecious shrub, were examined.
2 The objectives were to determine whether females allocate more biomass or nitrogen per reproductive episode than males, and whether there is a long-term cost of reproduction in terms of subsequent growth or reproduction for either sex. If there is no long-term cost of reproduction, then reproduction may be viewed as free in an evolutionary sense.
3 As is generally the case in dioecious species, females allocated more biomass and nitrogen to reproduction than males. Females also showed delayed costs of reproduction in terms of decreased growth and subsequent reproduction, whereas males did not.
4 The lack of measurable delayed costs in males suggests that with the evolution of dioecy, selection has reduced delayed costs of reproduction in S. grandiflora males. In contrast, females that were prevented from reproducing were able to re-allocate resources to growth, and produced more stem length on average than males. This re-allocation response may have evolved to reduce delayed costs of reproduction in females over time frames longer than that considered in the present study.     

Plant and insect genetic variation mediate the impact of arbuscular mycorrhizal fungi on a natural plant–herbivore interaction

1. While both arbuscular mycorrhizal (AM) fungi and plant and insect genotype are well known to influence plant and herbivore growth and performance, information is lacking on how these factors jointly influence the relationship between plants and their natural herbivores. 2. The aim of the present study was to investigate how a natural community of arbuscular mycorrhizal fungi affects the growth of the perennial herb Plantago lanceolata L. (Plantaginaceae), as well as its interaction with the Glanville fritillary butterfly [Melitaea cinxia L. (Nymphalidae)]. For this, a multifactorial experiment was conducted using plant lines originating from multiple plant populations in the Åland Islands, Finland, grown either with or without mycorrhizal fungi. For a subset of plant lines, the impact of mycorrhizal inoculation, plant line, and larval family on the performance of M. cinxia larvae were tested. 3. Arbuscular mycorrhizal inoculation did not have a consistently positive or negative impact on plant growth or herbivore performance. Instead, plant genetic variation mediated the impact of arbuscular mycorrhizal fungi on plant growth, and both plant genetic variation and herbivore genetic variation mediated the response of the herbivore. For both the plant and insect, the impact of the arbuscular mycorrhizal community ranged from mutualistic to antagonistic. Overall, the present findings illustrate that genetic variation in response to mycorrhizal fungi may play a key role in the ecology and evolution of plant–insect interactions.     

Sex determination and sex ratio in the dioecious shrub Salix viminalis L.

 Various ecological factors (e.g. herbivory, difference between males and females in colonising ability) have been invoked to explain female-biased sex ratios in populations of willow species. It was implicitly assumed that genetic factors would lead to a balanced sex ratio in the absence of ecological disturbances. In an experiment carried out in a homogeneous environment and in the absence of herbivores the progeny sex ratio of 13 crosses of basket willow (Salix viminalis L.) was observed to range from extreme female bias to extreme male bias. The observed sex ratio cannot be explained by the presence of sex chromosomes without assuming that additional loci are also involved in the sex determination. Alternatively, the sex ratios in this study can be explained by a sex determination mechanism governed by multiple independent loci. Received: 1 February 1996 / Accepted: 14 June 1996     

Variation of morphological and chemical traits in sexes of the dioecious perennial grass Poa ligularis in relation to shrub cover and aridity in Patagonian ecosystems

Sexes of dioecious species may have dimorphic responses to environmental variation due to differences in resource requirements and reproductive costs. We analyzed the effect of aridity/relative shrub cover, and vicinity to shrub patches on morpho-chemical traits of sexes of the dioecious perennial grass Poa ligularis in patchy arid ecosystems in northern-central Patagonia. We hypothesized that sexes of P. ligularis have dimorphic responses in morpho-chemical traits in relation to the environmental variation induced by aridity/relative shrub cover and vicinity to shrub patches. We selected seven sites across a gradient of increasing aridity and relative shrub cover. We randomly collected 5–10 P. ligularis plants per site registering the sex (female or male) and location with respect to shrub patches (shrub patch or inter-patch). For each plant, we assessed morpho-chemical traits (height of the vegetative tillers, length/dry weight/area of blades, specific blade area, nitrogen and soluble phenol concentration in blades). Sexes showed dimorphic responses in height of vegetative tillers, blade length, and blade area with respect to vicinity to shrub patches; and in variation of soluble phenolics in blades in relation to aridity/relative shrub cover. Responses in both sexes were opposite to those expected by aridity, highlighting the role of favorable environments induced by shrub canopies on dimorphic responses of sexes of P. ligularis. Resource-rich microsites associated with shrub canopies promoted increased plant performance of females with high reproductive costs while resource-poor open areas, favorable for pollen dispersal, induced improved chemical defenses of males. These results are consistent with the resource availability hypothesis.     

Floral display, pollinator visitation and reproductive success in the dioecious perennial herb Wurmbea dioica (Liliaceae)

Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998     

Breeding biology, resource partitioning and reproductive effort of a dioecious shrub, Clutia pulchella L. (Euphorbiaceae)

 The flowering of Clutia pulchella was studied in a coastal scarp forest community in southern KwaZulu-Natal, South Africa. Sex dependent differences in the frequency and magnitude of floral output were recorded, with male individuals producing 4.32 times more flowers per leaf axis than females. Increased node production, as a manifestation of significantly increased branching, allowed for prolific flower production in males. Energy investment per flower was marginally greater in females (0.045 J/mg), despite copious pollen production and glandular secretion in males (0.035 J/mg). Differential flower production, reproductive organ dry mass and an unbalanced branching ratio revealed that “per plant” expenditure towards flowering was more pronounced in males (♀ flowering × 5.04). The elevated cost of fruit production (♀ flowering × 5.5), inclusive of imperfect fruit set, resulted in overall reproductive expense being slightly higher for females. Such sex-dependent reproductive investment was shown to have minimal influence on the population sex ratio, with sex frequencies and nearest neighbor distances yielding a non-significant male bias of 1.16. No evidence of spatial asymmetry was found. Received December 1, 1999 Accepted September 27, 2000     

Root volatiles in plant–plant interactions II: Root volatiles alter root chemistry and plant–herbivore interactions of neighbouring plants

Volatile organic compounds (VOCs) emitted by plant roots can influence the germination and growth of neighbouring plants. However, little is known about the effects of root VOCs on plant–herbivore interactions of neighbouring plants. The spotted knapweed (Centaurea stoebe) constitutively releases high amounts of sesquiterpenes into the rhizosphere. Here, we examine the impact of C. stoebe root VOCs on the primary and secondary metabolites of sympatric Taraxacum officinale plants and the resulting plant‐mediated effects on a generalist root herbivore, the white grub Melolontha melolontha. We show that exposure of T. officinale to C.stoebe root VOCs does not affect the accumulation of defensive secondary metabolites but modulates carbohydrate and total protein levels in T. officinale roots. Furthermore, VOC exposure increases M. melolontha growth on T. officinale plants. Exposure of T. officinale to a major C. stoebe root VOC, the sesquiterpene (E)‐β‐caryophyllene, partially mimics the effect of the full root VOC blend on M. melolontha growth. Thus, releasing root VOCs can modify plant–herbivore interactions of neighbouring plants. The release of VOCs to increase the susceptibility of other plants may be a form of plant offense.     

Fine‐tuning of defences and counter‐defences in a specialised plant–herbivore system

1. The plant–herbivore arms race has been postulated to be a major driver for generating biological and biochemical diversity on Earth. Herbivore feeding is reduced by the production of chemical and physical barriers, but increases plant resistance against subsequent attack. Accordingly, specialisation is predicted to be an outcome of herbivores being able to circumvent plant‐induced defences. 2. Using a specialised plant–herbivore system, in which adult chrysomelid beetles (Chrysochus auratus) feed on leaves and larvae feed on roots of dogbane (Apocynum spp.), this study investigated whether root latex and cardenolides are effective against the soil‐dwelling larvae, and whether such defences could be circumvented by the herbivore. 3. Across two Apocynum species, C. auratus larvae were not affected by latex production or cardenolide amounts and diversity. By contrast, cardenolide apolarity was detrimental to larval growth. Yet larval feeding decreased average root cardenolide apolarity in A. cannabinum and larvae performed better on those plants. Finally, above‐ground induction rendered the plants more toxic by increasing root cardenolide apolarity and maintaining it, even during subsequent larval herbivory. 4. Therefore, the intimate relationship and interaction between Chrysochus and Apocynum are maintained by a delicate balance of herbivore manipulation and plant chemical induction.     

Proximity to canopy mediates changes in the defensive chemistry and herbivore loads of an understory tropical shrub,Piper kelleyi

Phytochemical traits are a key component of plant defense theory. Chemical ecology has been biased towards studying effects of individual metabolites even though effective plant defenses are comprised of diverse mixtures of metabolites. We tested the phytochemical landscape hypothesis, positing that trophic interactions are contingent upon their spatial location across a phytochemically diverse landscape. Specifically, intraspecific phytochemical changes associated with vertical strata in forests were hypothesised to affect herbivore communities of the neotropical shrub Piper kelleyi Tepe (Piperaceae). Using a field experiment, we found that phytochemical diversity increased with canopy height, and higher levels of phytochemical diversity located near the canopy were characterised by tradeoffs between photoactive and non‐photoactive biosynthetic pathways. For understory plants closer to the ground, phytochemical diversity increased as direct light transmittance decreased, and these plants were characterised by up to 37% reductions in herbivory. Our results suggest that intraspecific phytochemical diversity structures herbivore communities across the landscape, affecting total herbivory.     

The effects of drought and herbivory on plant–herbivore interactions across 16 soybean genotypes in a field experiment

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Male‐biased hermaphrodites in a gynodioecious shrub,Daphne jezoensis

Gynodioecy, a state where female and hermaphrodite plants coexist in populations, has been widely proposed an intermediate stage in the evolutionary pathway from hermaphroditism to dioecy. In the gynodioecy–dioecy pathway, hermaphrodites may gain most of their fitness through male function once females invade populations. To test this prediction, comprehensive studies on sex ratio variation across populations and reproductive characteristics of hermaphrodite and female phenotypes are necessary. This study examined the variation in sex ratio, sex expression, flower and fruit production and sexual dimorphism of morphological traits in a gynodioecious shrub, Daphne jezoensis, over multiple populations and years. Population sex ratio (hermaphrodite:female) was close to 1:1 or slightly hermaphrodite‐biased. Sex type of individual plants was largely fixed, but 15% of plants changed their sex during a 6‐year census. Hermaphrodite plants produced larger flowers and invested 2.5 times more resources in flower production than female plants, but they exhibited remarkably low fruit set (proportion of flowers setting fruits). Female plants produced six times more fruits than hermaphrodite plants. Low fruiting ability of hermaphrodite plants was retained even when hand‐pollination was performed. Fruit production of female plants was restricted by pollen limitation under natural conditions, irrespective of high potential fecundity, and this minimised the difference in resources allocated to reproduction between the sexes. Negative effects of previous flower and fruit production on current reproduction were not apparent in both sexes. This study suggests that gynodioecy in this species is functionally close to a dioecious mating system: smaller flower production with larger fruiting ability in female plants, and larger flower production with little fruiting ability in hermaphrodite plants.     

Indirect effects of cattle grazing on shrub spatial pattern in a mediterranean scrub community

Identifying the mechanisms and interactions that influence the spatial structure of vegetation is important for both scientific and practical purposes. Grazing is one of the most fundamental interactions in ecology but so far its effect on vegetation spatial pattern received little attention. In this study we propose a conceptual model that can be used to predict the effect of grazing on shrub spatial pattern in water-limited ecosystems where shrubs grow within a matrix of annual vegetation. According to the model, grazing may increase or decrease clumping in shrub distribution, depending on (1) the relative palatability of shrubs vs. annual plants to the herbivores, and (2) the manner (negative or positive) by which adult shrubs and annual plants affect the establishment of shrub seedlings. We tested our model in a Mediterranean scrub ecosystem by analyzing the development of shrub spatial pattern over a period of 40 years in plots characterized by contrasting intensities of cattle grazing. As predicted by the model, all plots showed a clumped pattern of shrub distribution in the absence of cattle grazing while intense cattle grazing reduced the clumpiness of the vegetation and generated a more random pattern of shrub distribution. Interestingly, plots representing the two grazing regimes did not differ significantly in their shrub cover, suggesting that shrub spatial pattern may be more sensitive to grazing than overall shrub cover.     

Gender-specific costs of reproduction on vegetative growth and physiological performance in the dioecious shrub Corema album

Background and Aims

Reproductive costs imply trade-offs in resource distribution at the physiological level, expressed as changes in future growth and/or reproduction. In dioecious species, females generally endure higher reproductive effort, although this is not necessarily expressed through higher somatic costs, as compensatory mechanisms may foster resource uptake during reproduction.

Methods

To assess effects of reproductive allocation on vegetative growth and physiological response in terms of costs and compensation mechanisms, a manipulative experiment of inflorescence bud removal was carried out in the sexually dimorphic species Corema album. Over two consecutive growing seasons, vegetative growth patterns, water status and photochemical efficiency were measured to evaluate gender-related differences.

Key Results

Suppression of reproductive allocation resulted in a direct reduction in somatic costs of reproduction, expressed through changes in growth variables and plant physiological status. Inflorescence bud removal was related to an increase in shoot elongation and water potential in male and female plants. The response to inflorescence bud removal showed gender-related differences that were related to the moment of maximum reproductive effort in each sexual form: flowering in males and fruiting in females. Delayed costs of reproduction were found in both water status and growth variables, showing gender-related differences in resource storage and use.

Conclusions

Results are consistent with the existence of a trade-off between reproductive and vegetative biomass, indicating that reproduction and growth depend on the same resource pool. Gender-related morphological and physiological differences arise as a response to different reproductive resource requirements. Delayed somatic costs provide evidence of gender-related differences in resource allocation and storage. Adaptive differences between genders in C. album may arise through the development of mechanisms which compensate for the cost of reproduction.